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VERZAMELD IN
NEDERLANDSCH OOST-INDIË 1899—19
00
AAN BOORD H. M. SIBOGA ONDER COMMANDO VAN
Luitenant ter zee T kl. G. F. TYDEMAN
UITGEGEVEN DOOR
Dr. MAX WEBER
Prof. in Amsterdam, Leider der Expeditie
(met medewerking van de Maatschappij ter bevordering van het Natuurkundig
Onderzoek der Nederlandsche Koloniën)
BOEKHANDEL EN DRUKKERIJ
VOORHEEN
E. J. BRILL
LEIDEN
Siboga-Exp editie
XXVIII a
THE
POLYZOA OF THE SIBOGA EXPEDITION
BY
SIDNEY F. HARMER, sc.a, f.r.s.
Keeper of Zoology in the British Museum (Natural History), London
PART I
ENTOPROCTA, CTENOSTOMATA and CYCLOSTOMATA
With 12 plates
=ê=-&©$-^=- —
late E. J. BRILL
PUBLISHERS AND PRINTERS
LEVDEN — 19 15
PART I
ENTOPROCTA, CTENOSTOMATA and
CYCLOSTOMATA.
CONTENTS.
I. IXTRODUCTION
II. DESCRIPTIONS OF SPECIES
Class POLYZOA J. V. Thomps. = BRYOZOA Ehrb.
Sub-Class I. ENTOPROCTA Nitsche . . .
page
I
Family Loxosomatidae Hincks 4
Loxocalyx Mort 6
1. Loxocalyx lincatus n. sp 6
2. „ leptoclini Harm 8
Loxosoma Kef. 8
Loxosoma lanchesteri n. sp 8
n sluiteri n. sp 9
„ annulatitm d. sp II
„ velatum n. sp 13
„ cirriferum n. sp 14
„ circulare n. sp 16
„ pitsillum n. sp 16
„ spp. incert. (on Retepora) 17
9-
10.
11.
12.
Loxosoma troglodytts n. sp 17
„ 0;"Mte II. sp 19
v subsessiie 11. sp 19
„ loricatum n. sp 21
„ coccifortne n. sp 22
Family Pedicelunidae Johnst 23
Pedicellina M. Sars 23
Pedicellina compacta n. sp 24
Barentsia Hincks 25
Barentsia gracilis M. Savs 27
n disercta Busk 29
„ /.Mi; Kirkp 32
„ gcniiitlata n. sp 33
Sub-Class II. ECTOPROCTA Nitsche 35
Order I. GYMNOLAEMATA Allm 35
Sub-Order I. CTENOSTOMATA Busk 35
Group A. Carnosa Gray 35
Family Flustrellidae Hincks 38
Elzerina Lamx 38
1. Elzerina blainvillii Lamx 38
Family Alcyonidiidae Johnst 36
Alcyonidium Lamx 36
AlcyoniJium polyoum Hass 37
PALUDICELLEA, STOLONIFERA and VESICULARINA 41
Group B. Paludicellea Allm 43
Arachnoidea J. E. S. Moore 50
1. Arachnoidea pro/cc/a n. sp 5°
Family Nolellidae nom. nov 52
Nolella Gosse 52
1. Nolella papuensts Busk 53
2. „ annectens n. sp 57
Family Yictorellidae Hincks 44
Victorella ïav. Kent 44
Victorella sibogac n. sp 45
Family ARACHNIDIIDAE Hincks 48
Arachnidium Hincks 48
Arachnidium irrcgularc n. sp 49
,482J
VI
CONTENTS.
Family Vesicularhdae John t.
Vesicularia J. V. Thomps.
u/aria fapuensis Busk
Amathia Lams . . .
Amathia convoluta Lamx
Group C. Vesicularina Johnst.
.... 60
61
61
64
64
2. Amathia distans Busk
Bowerbankia Farre
I. Bowerbankia imbricata Adams (? sp.) .
Group D.
Farn. VALKERIIDAE Hincks
Valkeria Flem
Valkeria atlantica Busk
v tuberosa Heil
Family MlMOSELLIDAE Hincks
Mimosella Hincks
Mimosclla bigeminata Wateis
n verticillata Heil
3. „ tettuis n. sp
Distribution of Extoprocta and Ctenostomata . . .
Stolonifera Ehlers
1.
2.
1.
2.
73
73
73
76
78
78
79
Si
84
Family Buskiidae Hincks . .
Buskia Alder
1. Buskia nitens Alder
2. „ seiigera Hincks
3. „ pilosa n. sp
(Family Triticellidae G. O. Sars
Triticella Dalyell . . . .
I. Triticella boeckii G. O. Sars
Sub-Order II.
Family Crisiidae Johnst. .
Crisia Lamx ....
Crisia elongata Milne Edw
„ cuneata Mapl
„ ierguelemis Busk
„ geniculata Milne Edw
Family ENTALOFHORIDAE Reuss
Entalophora Lamx . .
Entalophora proboscidea Milne Edw. . . .
„ delicatula Busk
„ intricaria Busk (? sp.) ....
Family DlASTOPORIDAE Busk .
Berenicea Lamx . . .
Berenicea sarniensis Norm
lineata MacGill
CYCLOSTOMATA Busk
96
96
96
Family TUBULIPORIDAE Johnst.
Reptotubigera D'Orb, . .
1 . Reptotubigera philippsae n. sp
Tubulipora Lamx . . .
1. Tubulipora concinna MacGill
2. „ atlantica (Forbes, MSS.) Johnst.
■03
105
106
107
107
10S
110
112
"3
114
114
116
118
119
120
122
123
124
2<7. Tubulipora atlantica (Forbes, MSS.) Johnst., \3.x.Jiexuosa
1'ourt
3. „ pulcherrima Kirkp
4. „ cassiformis n. sp
Crisina D'Oib. . . .
I. Crisina radio ns Lamk
Tervia Juli
I. Tervia jellyae n. sp
Specimens in Storna lopora-condiüon
Family Horneridae Smitt .
Hornera Lamx . . .
1. Hornera spinigera Kirkp
2. „ caespitosa Busk
Family Cytisidae D'Orb. .
Supercytis D'Orb. . . .
1. Supercytis watersi n. sp
Family Lichenoporidae Smitt
Lichenopora Defrance . .
1. Lichenopora novae-zelandiae Busk
2. „ buski n. sp
3. „ mediterranen Mich. (? sp.) . .
Distribution of Cvclostomata .
LJBI.IOGRAPHY
INDEX
EXPLANATION OF THE 1'I.ATES.
page
60
68
70
70
72
85
85
85
87
89
90
90
90)
92
96
127
129
135
137
139
143
143
144
147
147
147
149
150
150
151
153
153
155
161
164
166
171
173
I. INTRODUCTION.
It has usually been supposed that the conditions under which marine animals occnr in
the Tropics are specially unfavourable for the growth of Polyzoa. Lamouroux *), speaking of
"Corallines", states that "il semble qu'une grande chaleur leur soit contraire"; while Ortmann 2)
remarks "Anderseits scheinen die Tropen eine Grenze für die Bryozoengebiete zu bilden"; and,
ten years later, Nordgaard 3) accepts the view that Polyzoa are rare in low latitudes. It has
indeed been clear for some time that the belief in the absence of Polyzoa from Tropical localities
is due merely to insufficiënt knowledge; but if any further proof were needed, the results of the
'Siboga' Expedition would be amply sufficiënt to disprove the view that Tropical conditions are
in any way antagonistic to the growth of Polyzoa. It may safely be asserted that, in Malay
waters at least, Polyzoa are present in large numbers, both of species and of individuals; and
that they occur from the littoral region down to the greatest depths investigated. The 'Siboga'
collection is far the richest that has at present been made in any Tropical region ; and it is
specially important in supplying information with regard to a district which had previously been
almost unknown. Further to the South, the Australian forms have received much attention
from Busk, MacGili.ivray, Hincks, Waters, Maplestone and others. Ortmann's paper, already
referred to, gives some information with regard to Japanese species. But with the exception of
papers by Hincks 4), Kirkpatrick 5), Miss Thornelv i!) and Waters 7), little recent work has
been done on the marine Polyzoa of the Indo-Malay region, the importance of which from the
point of view of Geographical Distribution need not be emphasized.
The official list of the localities at which specimens were collected during the voyage ot
the 'Siboga' includes 323 numbered Stations; and Polyzoa were found in about 134 of these.
In some cases the number of species obtained was very large. Thus in a single bottle from
Station 144 about 38 species were discovered. Although this is very much more than the
average number found, the process of sorting the material has been a very arduous one,
1) Lamouroux, J. V. F., 1816, p. xxxm.
2) Ortmann, A., 1889, p. 67.
3) Nordgaard, O., 1900, "Norske Nordhavs-Exp.", XXVII, "Polyzoa", p. 27.
4) Hincks, T., 1884, 1887.
5) Kirkpatrick, R., 1888, 18901, 1S902.
6) Thornely, I.. R., 1905, 1907, 1912.
7) Waters, A. W., 1909, 1910, 1913, 1914.
1
SIP.OG A-EXPEDITIE XXVIII a.
particularly since it has been necessary to subject nearly the whole of it to a careful micros-
copical scrutiny. In a large number of cases a specimen which was placed in the collection oi
Polyzoa, during the original sorting of the material, has been found, on examination, to have
other species growing on it; and many of the most interesting specimens have been discovered
as the result of their occurrence on some other species which was conspicuous enough to be
referred to the Class under consideration. I cannot doubt that if it had been possible to examine
the entire collection, of animals of all kinds, many Polyzoa would have been recorded which
have escaped discovery. This is particularly likely to be true of the more inconspicuous forms,
such as Entoprocta and Ctenostomata, for instance. The genus Loxosoma, belonging to the
former group, is obviously very abundant in Malay waters. In other parts of the world, species
of Loxosoma have been found commonly on such animals as Sponges, Tunicates, Polychaets
and Gephyrea. Most of the specimens of Loxosoma which presumably occurred on members of
these groups collected by the 'Siboga' have escaped detection, as the "hosts" in question were
not submitted to me for examination. Discoveries of specimens of Loxosoma on members of
these groups have for the most part been due to the accidental inclusion of fragments of the
respective "hosts" in bottles labelled as Polyzoa. The systematic examination of the 'Siboga'
collection of Hydroids would almost certainly have resulted in a considerable addition to the
list of specimens and species recorded; while it may further be pointed out that the material
submitted for examination included hardly any Algae.
The collection worked out was received in 583 bottles and tubes, which were numbered
consecutively from 1 to 583 in the order in which they happen to have been received. Although
these numbers were given, in the first instance, for convenience of reference during the progress
of the work, I have considered it important to record them in the subjoined systematic account
of the collection. It will thus be possible to yerify or correct my statements by reference to
the actual specimens on which they are respectively based. I think it well to emphasize this
point because, in some accounts which have been published — for instance in Busk's 'Challenger'
Reports — it is sometimes difficult to know what specimens are referreel to, in the case of an
account based on specimens from several distinct localities, and perhaps not rightly referred to
the same species.
The specimens here recorded thus bear arbitrary numbers, from 1 to 583; and in each
case the species found in a single bottle are further distinguished by letters (A, B, C ),
given to them in the order in which they were sorted out. Every specimen in the collection
thus bears a distinctive symbol of its own. I propose to give a list of the specimens examined,
arranged under Stations, at the end of the Report.
In view of the fact that no general account of the Oriental species has hitherto been
published, and of the convenience of having the description of the Polyzoon-fauna of so important
a district recorded in one place, I have ventured to prepare figures of nearly all the species,
even though many of them are already well knovvn. This course appears to me the more
defensible because the publication of figures will enable other workers to criticise more effectively
the correctness of my determinations. In many works which have been published — and on
which, it may be, conclusions relating to Geographical Distribution have been based — the
reference of a specimen to a particular species depends entirely on the unsupported statement
of the author of the Memoir in question ; and it cannot be doubted that in a considerable
proportion of cases the determinations have been incorrect.
In addition to the specimens collected by the 'Siboga' I have made special use of the
following collections :
(i) A collection made by Dr A. C. Haddon in Torres Straits, 1888 — 1889. This formed
the subject of a paper by Kirkpatrick j), and the specimens thus described are preserved in
the British Museum. A supplementary part of the Collection, which had not been examined, was
subsequently presented by Professor Haddon to the University Museum of Zoology, Cambridge
(Reg. Feb. 24, 1898), and was found to contain a number of species which had not been
recorded by Kirkpatrick. The specimens in the Cambridge collection are the ones which are
more particularly considered in the present Report.
(2) Specimens from Singapore, presented to the Cambridge Museum by the late Professor
C. Stewart (Reg. April 26, 1899); the late Mr F. P. Bedford (Reg. Nov. 11, 1899); and
Dr R. Hanitsch (Reg. June 19, 1900).
(3) A collection from Japan, in the neighbourhood of Tokyo, received at the Cambridge
Museum from Mr A. Owston (Reg. June 23, 1902). These are included only so far as they
appear to belong to the same species as those contained in the other Collections.
1) Kirkpatrick, R., 1S90-.
II. DESCRIPTIONS OF SPECIES.
/
Class POLYZOA J. V. Thompson, 1830.
= BRYOZOA Ehrenberg, 1831.
Sub-Class I. ENTOPROCTA1) Nitsche.
1869, Zeitschr. wiss. Zool., XX, p. 596.
Fam. Loxosomatidae 2) Hincks.
Loxosomidae Hincks.
18S0, "Hist. Brit. Mar. Pol.", p. 571.
I have indicated above my reasons for believing that the species of this Family here
recorded form but a small proportion of those that really occur in Malay waters. The specimens
found have been discovered, without exception, accidentally, in the course of the examination
of some other species. The Family is obviously abundantly represented in the Indo-Malay Archi-
pelago ; and it cannot be doubted that a more systematic search would result in the discovery of
other species, particularly if attention were paid to Sponges, Polychaets, Gephyrea and Tunicates.
I am acquainted with but few references to the Loxosomatidae outside the Atlantic,
Mediterranean and Arctic areas; and it may be worth while to mention the cases in question.
In a paper on the Hydrozoa and Polyzoa of the China Sea, Kirkpatrick 8) has recorded L.
crassicaudq.} from the Tizard Bank, 27 fathoms. After examining the original material (Brit.
Mus., 89. 8. 21. 71.) I am unable to refer any of the 'Siboga' specimens to the same species;
and the condition of Mr Kirkpatrick's specimens does not justify me in expressing an opinion
with regard to the correctness of his determination. Whitelegge 4) has recorded Loxosoma sp.
from Phascolosoma australis (sic). Haswell b) has mentioned an undetermined Loxosoma which
he found on the large chaetae of Coppingcria longisctosa at Port Molle, Oueensland. In his
1) By many writers tlie name ENDOPROCTA is employed. I know of no reason for departing from the form originally
introduced by Nitsche.
2) It was recognized by KowalewsKY (1S66. "Beitr. Anat. u. Entw. Loxosoma neapolitanum" ', Mem. Acad. Imp. St Pétersbourg,
7; X, N° 2), that Loxosoma must lic regarded as the representative of a new Family at least.
3) Kirkpatrick, R., 18901, p. 17.
4) WHITELEGGE, T., 1890, "List Mar. & Fresh-water Inv. Kanna Port Jackson", J. Proc. Roy. Soc. N. S. Wales, XXIII, Pt II, p. 293.
5) 11aswi.ii. \V. A., 1891, "Obs. Chlovaemidae", Proc. Linn. Soc. N. S. Wales, (2) VI, p. 330, PI. XXVI, rig. 1.
4
5
Report on the Polyzoa of the Belgian Antarctic Expedition, Waters l) states that he has found
a Loxosoma (undetermined) among material from Lizard Island, Oueensland; and the same
author -) has more recently recorded L. kcfersteinii from the Sudanese Red Sea. I have myself
found a species (probably new) of Loxosomatidae, in the Collection of the British Museum
(86. 6. 8. 3.), growing abundantly on Ainathia wilsoni Kirkpatrick, from Port Phillip, Victoria.
I am not convinced that Calman's record3) of Loxosoma'*. on a Crab [Xanthias haswelli) from
Christmas Island, Indian Ocean, really refers to this genus.
The discrimination of the species of Loxosoma is often very difficult in the absence of
living specimens, or at least of specimens which have not been killed with special care. The
number of the tentacles is a character of importance in clistinguishing the species from one
another ; and unless the animals have been killed in an extended condition it cannot always be
ascertained with certainty. I am aware of the fact that my descriptions leave much to be desired.
The following is a list of the species of Loxosomatidae which I have distinguished in
the present Report; with the approximate number of their tentacles and the average length of
fully adult specimens. It must, however, be remembered that the state of preservation is not
good enough to allow me to speak with certainty as to the number of tentacles in some cases;
so that a margin of error must be allowed for in the third column. The total lenaths oiven
(calyx -j- stalk) are at most an indication of the relative sizes. It may be presumed that in most,
if not all, cases the length of the living animal, with fully extended stalk, materially exceeds
these measurements.
SPECIES.
"HOST'
Approximate
number
of tentacles
Total length,
Loxocalyx
Loxocalyx
Loxosoma
Loxosoma
Loxosoma
Loxosoma
Loxosoma
Loxosoma
Loxosoma
Loxosoma
Loxosoma
Loxoscma
Loxosoma
Loxosoma
lineatus 11. sp.
leptoclini Harmer
lanchesteri n. sp.
sluiteri n. sp.
annulatum n. sp.
velatum n. sp.
cirriferum n. sp.
circulare 11. sp.
pusillum n. sp.
trogtodytes 11. sp.
breve n. sp.
subsessile 11. sp.
loricatum n. sp.
cocciformc n. sp.
Halicliondria.
Cephalodiscus sibogae Harmer
Pliascolosoma pellucidum var.
Pliascolwn convestitus Sluit.
Retepora, Sclüzoporclla.
Retepora.
Retepora.
Retepora.
Retepora.
Lepralia celleporoides Busk.
Schizoporella, Ccllepora.
Conescliarellina.
Ca n da.
Siphonicytara.
10?
900
10
500
20 +
1230
8
400
9
500 — 600
14 — 16
500
14— iS
650
12
300
8—10
250
9
190
9
200
8
120
8(10)
180
10 — 12
260
5*
In a recent work Mortensen 4) has proposed the division of Loxosoma into three genera,
which he characterises as follows :
0 Waters, A. W., 1904, p. 100.
2) Ibid., 1910, p. 252.
3) Calman. W. T., 191 1, Ann. Mag. Nat. Hist. (8) VIII, p. 550.
4) Mortessex, Th., 191 1, "A new Species of Entoprocta, Loxosomelta antedoms", Danmark-Exspeditionen til Grcnlands Noid-
Ostkyst 1906 — 1908, Bind V, Xr 8, Kobenhavn.
(I) Loxosoma Keferstein : Genotype L. singulare Keferstein.
The foot is a sucking disc, provided with straight and oblique muscles. No foot-
gland, even in the buds.
(II) Loxocalyx Mort. (nov.) : Genotype L. raja Schmidt.
Foot-gland present in the adult, as well as in the buds; the foot usually with
wing-like expansions. Only straight muscles are developed in the foot.
(III) Loxosomclla Mort. (nov.): Genotype L. crassicauda Salensky.
Foot-gland present in the buds, but not in the adults. Only straight muscles are
present in the foot.
The persistence of the foot-gland in the adult is a feature which marks off a certain
number of the species from the rest ; and perhaps entitles them to genene rank. I feel less
convinced that the other species are sufhciently well known to warrant the separation of
Loxosomella from Loxosoma.
Loxocalyx Mortensen.
191 1, t. cit., p. 406.
The 'Siboga' collection contains two species which are referable to this genus. The state
of preservation of the first of these is not good enough to allow several important points to
be made out. I am unable, for instance, to state definitely what is the number of the tentacles;
but on the whole I think I am justified in describing it as a new species.
I therefore record it under the name of
1. Loxocalyx lineatus n. sp. (PI. II, figs. 1 — 3).
332. C, on a Siliceous Sponge [Halichondria sp. ')). Stat. 64. Kambaragi Bay, Tanah Djampeah,
o — 32 Metres; coral, coral-sand.
Calyx from 220 to 320 u. long, 190 — 230 y. broad; with moderately developed lateral
membranous expansions. Number of tentacles uncertain, perhaps 10. A series of vesicular cells
encircles the distal part of the lophophore. Buds not numerous. Stomach with well developed
lateral lobes. Stalk from 570 to 670 y. long, usually about twice the length of the calyx ;
marked by four longitudinal lines (the nature of which is uncertain), to which the specific name
refers. Foot with well marked alate expansions and a persistent foot-gland.
The specimens were found on the outside of a species of Halichondria, of narrow
cylindrical form.
Several Polychaets belonging to the family Syllidae occurred in the interior of the
cylindrical branches of the Sponge.
The examples selected for figuring bear only one bud each, represented merely by a
subspherical swelling. In other individuals the bud of one side had nearly reached its limit of
1) I am indebted to Mr R. KlRKI'ATRlCK for tliis determination.
6
growth, but the next bud of the same side was merely a minute tubercle. It may accordingly
be concluded that in this species there is only one vvell developed bud at a time.
The following species of Loxosoma are referable to Mortensen's genus Loxocalyx; though
the specific characters are by no means well ascertained in all cases; and it is uncertain whether
they all deserve specific rank :
Loxosoma alatum Barrois 1877.
Loxosoma cochlear Schmidt, 1876.
Loxosoma leptoclini Harmer, 1885.
Loxosoma ncapolitanum Kowalevsky, 1866.
Loxosoma pes Schmidt, 1878 (which has been regarded as a synonym of L. alatum).
Loxosoma ra ja Schmidt, 1876.
Loxosoma tcthyae Salensky, 1877.
Of these, L. alatum, L. cochlear, L. pes, L. raja and L. tetJiyae have been recorded
as occurring on various Sponges. In most of these species the stalk appears to be less than
twice the length of the calyx, but in L. tcthyae it is generally at least twice as long. In this
respect the 'Siboga' specimens agree more nearly with that species than with any of the others.
No adequate figure of L. tcthyae appears to have been published; but it is worth calling
attention to certain figures, by O. G. Costa, which have not, I think, been previously cited in
the accounts of this species, but which are of interest as being probably the earliest figures of
any species of Loxosoma. The figures in question were published in Cosïa's "Fauna del Regno
di Napoli", "Zoofiti", the title-page of which bears the date 1838. The real dates of publication
are given in 'Isis', 1846, Heft IX, p. 718; and relying on the information there given and on
the dates which occur on the edges of the sheets, the Section headed "Genere Tezia; Tethya
Lamk." was really dated Nov. 18, 1843 (pp. 1 — 8), Jan. 3, 1844 (pp. 9 — 16), and May 14, 1844
(pp. 17 — 24). On pp. 11 and 12 of this Memoir Costa gives an account of certain polyp-like
structures which he regarded as belonging to the Tethya. The figures 1, 2 and 5, at the top
of PI. I, illustrating the description, are perfectly characteristic representations of Loxosoma
tcthyae-, figs 1 and 5 showing the general form of the calyx and stalk, and fig. 2 being a
good representation of the foot, and showing further the arrangement of the ectoderm-cells of
the stalk in eight longitudinal rows; a feature to which I called attention in my Memoir on
Loxosoma l), without being at that time acquainted with Costa's figures and description. It need
hardly be remarked that as Costa regarded the Loxosoma as a part of the Sponge, he did
not give it any generic or specific name.
The calyx of L. tcthyae, as seen in Costa's figures, is relatively narrow, and is not
furnished with thin lateral expansions. There can thus be little doubt that the 'Siboga' specimens
are not to be referred to that species. From L. leptoclini they differ in the absence of the
group of cells, at the distal end of the calyx, characteristic of that species 3), as well as by the
much longer stalk. L. raja Schmidt3) has a specially broad calyx; but its broadest part is at
1) Harmer, S. F., 1885, Quart. J. Mier. Sci., XXV, p. 262.
2) Ibid., PI. XIX, fig. 2.
3) Schmidt, O., 1876, Arch. mikr. Anat., XII, PI. I, fig. 1.
7
8
its proximal end. L. cocJilcar Schmidt x) has only 8 tentacles, a number which is almost certainly
smaller than that of the 'Siboga' specimens. L. pcs ~) has a stalk which is commonly shorter
than the calyx. L. neapolitanum '"') is very insufhciently known ; but its calyx is narrow and the
stomach does not appear to have vvell marked lateral lobes. L. alatum was very incompletely
characterised by Barrois 4), its original describer, and it is almost impossible to decide whether
this form is identical with any of the others or not. The specimens which have more recently
been referred to this species by Jullien and Calvet 5) are very different from Barrois' figure
in the proportions of the stalk, which in Jullien and Calvet's specimens is considerably shorter
than the calyx; and it appears doubtful whether they belong to L. alatum.
It seems justifiable, as the result of these considerations, to regard the 'Siboga' specimens
as the representatives of a new species.
2. Loxocalyx leptoclini Harmer. (PI. I, fig. i).
Loxosoma leptoclini Harmer, 1885, Quart. J. Mier. Sci. XXV, p. 263.
402. D., on Cephalodiscus sibogae (slide 402. A.). Stat. 204. Between Islands of Wowoni and
Buton, 75 — 94 Metres; sand with dead shells.
A single specimen of Loxocalyx was found on one of the slides of the above species
of Cephalodiscus. The foot-gland is well developed and the foot has strongly marked alate
expansions. The calyx is hardly more than half as long as that of L. lineatus\ and the stalk
is not quite twice as long as the calyx. The lateral lobes of the stomach are strongly marked,
and there are almost certainly ten tentacles. The calyx measures 180 y. in length, and 147 a
in breadth. The stalk is 320 u. long.
The specimen differs so much in size from those described above as L. lineatus that it
can hardly be referred to the same species. The brood-pouch contains two embryos, and ovaries
are present, without testes. Its measurements approach closely those of L. leptoclini, with which
it agrees in the general proportions of the stalk and body, in the number of the tentacles, in
the prominence of the lateral lobes of the stomach, and in the form of the wings of the foot.
It appears to me better to refer it provisionally to this species than to describe it as new.
Loxosoma Keferstein.
1862, "Unt. nied. Seethiere", Zeitsch. wiss. Zool. XII, p. 131.
1. Loxosoma lanchesteri n. sp. (PI. II, figs 4, 5).
Size large. Calyx pear-shaped, narrower distally, the broadest part close to the proximal
end and almost equalling its length. Lateral parts of the calyx somewhat depressed, the median
part more convex. Stalk much longer than the calyx, cylindrical proximally, and without foot-
1) In Brf.hm's "Thierleben", Ed. 2, 1878, Bd X, fig. on p. 1S1.
2) Cf. my remarks on this species, t. cit., p. 262.
3) Kowai.evsky, A., 1866, Mém. Acad. St Péteisbourg, (7) X, N° 2, figs I, 2.
4) BARROIS, J., 1877, "Rech. Emb. des Bryozoaires", p. 9, PI. XVI, fig. 4.
5) Jullien, J. and Calvet, L., 1903, "Rés. Camp. Sci. Prince de Monaco", XXIII, p. 29, PI. II, figs 40—4^.
8
gland in the adult, but dilating as it approaches the calyx, which, like the distal part of the
stalk, is marked by transversely running grooves, giving a crenulated appearance to the profile
of these parts. Stomach vvith some indication of lateral lobes. Buds as many as four at a time
(two on each side). Tentacles numerous, at least 20 in number. Sexes apparently separate.
The specimens to which this description refers do not form part of the 'Siboga' Collection,
but were found at Singapore and Pulu Java, Malacca, by Mr W. F. Lanchester, after whom
I have the pleasure of naming the species, on the tails of two specimens of a Sipunculid,
determined by him as Phascolosoma pellucidum var. They were presented to the University
Museum of Zoology at Cambridge, but the type-specimen will be transferred to the British
Museum, while a duplicate will be placed with the 'Siboga' Collection.
The material is unfortunately badly preserved, and some of it was further injured by the
use of oil of cloves, which produced great contraction. The specimens transferred from absolute
alcohol directly to Canada balsam dissolved in the same fluid l) are much more satisfactory.
The measurements of the largest specimen found (fig. 4) are as follows : — length of
calyx, 346 'j.\ breadth of calyx, at its widest part, 308 fi; length of stalk, 896 y.\ breadth of
stalk, at the proximal end, 58 y. ; breadth of the stalk, just before it joins the calyx, 134 u..
The other specimens are smaller-, the smallest having the measurements (given in the same
order) 180, 166, 300, 51 and yj; two other individuals being intermediate in these measurements
between the largest and the smallest specimens. It will be seen that the individuals of the
present species may be described as very large ; and this is sufficiently brought out by comparing
the figures given with those of other species 2).
The number of tentacles cannot be given with certainty; but it is at least 20, and is
probably more. The alimentary canal is not well shown, but the intestine, rectum and some
indication of the stomach may be seen in fig. 4. The specimen here represented appears to be
a male; a pair of organs which are almost certainly testes being- visible in the figure. The buds
are only visible in one of the slides which is not successfully mounted; but, in one individual,
four buds are present; — a large one on each side (the two not differing much in size) and
a small one at the base of each of the larger ones. The buds are not well enough preserved
to decide the question whether a food-gland is present or not.
It can hardly be doubted that the present species is different from other species which
have hitherto been described from the tails of Sipunculids [L. fihascolosomatum Vogt, 1876;
L. murmanicum Nilus, 1909; L. brumpti Nilus, 1909; L. minutum Osburn, 1910; and L.
sluitcri, described below). It has some resemblance to L. antedonis, described by Mortensen
(191 1) from the cirri of Hathrometra (Antedon) prolixa Sladen, from Greenland.
2. Loxosoma sluiten' n. sp. (PI. I, fig. 7).
373. A. On Phascolion convestitus Sluiter. Stat. 105. 6° 8' N., i2i°i9'E., 275 Metres; coral-bottom.
Calyx about 150 u. in length, and about 85 a in breadth, without cirri or lateral
1) For the use and advantages of this method, see my paper in Quart. J. .Mier. Sci., XI. VI, Pt 2, 1902, p. 264.
2) The figures were all drawn to the same scale ; but it must be noted that while those on PI. I were only reduced 2/3, those
on PI. II are reduced '/2.
S1BOGA-EXPEDITIE XXVIII O.
io
expansions. Buds? Tentacles 8. Stomach without lateral lobes. Stalk from one and a half to
twice as long as the calyx, reaching a Iength of 310^., ending in a small disc of attachment,
without food-gland.
The specimens here described were sent to me, in June, 1902, by Prof. C. Ph. Sluiter,
who had found them in the course of his examination of Phascolion convestitus, which has been
described by him as a new species in the 'Siboga' Report l) on the Sipunculids and Echiurids
obtained by the Expedition. I have the pleasure of naming this species of Loxosoma after
its discoveren
The occurrence of the Loxosoma has already been recorded by Prof. Sluiter in his
Report. As there pointed out, the Polyzoon occurred on all the specimens of the Phascolion,
principally on the posterior extremity of the body, although a few were found on the anterior
end as well.
Loxosoma has frequently been recorded on the skin of Sipunculids. The best known
species is L. phascolosomatum Vogt, usually found on species of Phascolosoma ; differing in man)'
respects from the 'Siboga' form, and notably in the great Iength of its stalk. A Loxosoma was
recorded by Brumpt ~) from Pliascolion strombi at Roscoff ; but it was neither described nor named.
Two species have more recently been described by Nilus 3) from Phascolion spitzbergensc found
in the Kola Fjord, on the Murman Coast of Barents Sea. Of these, L. murmanicum was found
on the anterior end of the Pliascolion, and L. brumpti on its posterior end. The aboral ends
of the calyx and the stalk of L. murmanicum are covered by a thick brown cuticle; thus
differing from the 'Siboga' species. L. brumpti does not possess this thick cuticle, but bears
two projecting organs, believed to be sense-organs, one on either side of the oral end of the
calyx. Both the species described by Nilus are said to have 8 tentacles; but it may be noted
that in fig. 8, given by that author of L. brumpti, 9 are indicated. L. minutum Osburn 4), found
on Pliascolion strombi and Phascolosoma cremita in the Woods Hole region, E. coast of N.
America, is less than "5 mm. in Iength, and has "about eight" tentacles. The stalk is shorter
than the calyx and terminates in a well marked, circular pedal disc.
I have had some hesitation in giving a new name to the 'Siboga' species, since I have
been unable to obtain any evidence with regard to some of its most important features, such
as the number of the buds and the character of the gonads. The absence of buds is specially
noteworthy, particularly in view of the fact that many of the individuals are small, with a
calyx-length of not more than 76 u.. It is possible that they are all young individuals which
have been produced by the metamorphosis of larvae. The 'Siboga' species is a very small form,
its calyx-length of 150 u. being much smaller than that of L. murmanicum (352 p.), which,
moreover, has a very short stalk, or than that of L. brumpti (320 u.), which it more nearly
approaches. The number of tentacles (8) is identical with that of both species described by Nilus.
So far as I have been able to ascertain, there is 110 specific difference between the
1) Monogr. XXV, 1902, p. 32, 33.
2) Brumpt, Ë., 1807. Arch. Zool. Exp. (3) V, p. 494.
3) Nilus, G., 1909, Trudui StPeterb. Obschch. XI, I, N° 4, c. r. Séances, p. 157.
4) Osburn, R. ('., 1912, Buil. l'.ur. Fisheries (Washington, I>. C). XXX, p. 212.
10
I I
individuals which occur at the anterior end of the Phascolion and those which are found on
its posterior end.
It may be useful to note a few other references to the occurrence of Loxosoma on
Sipunculids, and particularly on Phascolion. Besides the records of L. phascolosomatum on
Phascolosoma the following may be mentioned :
L. phascolosomatum , found on Phascolion by Andersson J) in E. Greenland and by
Norman '2) in E. Finmark.
Loxosoma sp., found on Phascolion by Marion3) at Marseilles; by Levinsen4) in the
Kara Sea: and by Théel 5) in the Gullmarfjord, S.W. Sweden : on Phascolosoma by Verrill i;)
on the Atlantic Coast of the United States ; and on Phascolosoma australe by Whitelegge 7)
in Port Jackson.
3. Loxosoma anuulatum n. sp. (PI. I, ligs 2 — 6).
Type. 195. D. On Retepora, 195. A., Stat. 274. 50 28.2 S., 1340 53.9 E., 57 Metres; sand and shells,
stones.
24. E. On Retepora, 24. A., Stat. 53. Bay of Nangamessi, Sumba, o — 36 Metres; coral-sand ;
near the shore, mud. (On Mimosella tennis slides, 24. C.2, 24. C.3).
133. H. On Retepora, 133. B., Stat. 164. I°42'.S S., I30°47'.5 E., 32 Metres; sand, small stones
and shells.
247. E. On Retepora, 247. A., Stat. 310. 8° 30' S., U9°7'.5E., 73 Metres; sand and dead coral.
[For other specimens which appear to belong to this species, see below, p. 13].
Calyx rather small relatively to the stalk, usually from 140 to 190 \i. long and 100 to
1 30 u. broad ; somewhat broader distally than proximally, where it passes continuously into the
stalk, without a sudden diminution in breadth. Cirriform organs small, usually one pair, opposite
the distal end of the stomach (fig. 2) ; occasionally one or two cirri on the more distal part
of the calyx. Tentacles 9, with an unpaired distal tentacle ; rarely 11. Stomach without lateral
lobes. Gonads ? Buds usually on one side of the calyx only, not more than two at a time, the
tip of the stalk prolonged beyond the attachment to the parent in the old state. Stalk varying
greatly in length, but always longer than the calyx, sometimes more than three times as long;
broader, relatively to the calyx, than in other species found on Retepora, conspicuously annu-
lated, without foot-gland, which is, however, present in the buds.
The specific name refers to the annulated character of the stalk : — a feature which
may be due to contraction, but is not seen in the other species, similarly preserved.
I am not acquainted with previous records8) of the occurrence of Loxosoma on Retepora \
but, judging from the evidence of the 'Siboga' Collection, it must be extremely common on
1) Andersson, K. A., 1902, "Bry. schwed. Exp.", Zool. Jahrb. Syst., XVI. p. 555.
2) Norman. A. M., 1903, "Notes Nat. Hist. E. Finmark", Ann. Mag. Nat. Hist. (7) XI, p. 574.
3) Marion, A. F., 1879. "Drag. Marseille", Ann. Sci. Nat., Zool. (6) VIII, p. 33.
4) Levinsen, G. M. R.. 1S86, "Bry. Kara-Havet", Dijmphna-Togtets zool.-bot. Udb., p. 327.
5) Théel, H., 1907, "Sver. Zool. Hafsstatiou Kristineberg", Ark. f. Zool.. IV. p. 71.
6) Verrill, A. E., 1879, "Prelim. Check-List Mar. Inv.", Prepared for the U.S. Comm. of Fish and Fisheries, Author's Edition,
New Haven, p. 31.
7) Whitelegge, T., 1890, "List Mar. and Fresh-water Fauna Port Jackson", J. Proc. Roy. Soc. N. S. Wales, XXIII, p. 131.
8) Except my own preliminary notice, referred to on p. 12.
11
I 2
species of that genus in Malay waters. Specimens of Retepora which I refer provisionally to
seven species have been found to be thus infested. In many cases, the Loxosoma was only
discovered during the examination of a microscopic preparation of the Retepore ; and it cannot
be doubted that I have failed to detect it on many specimens on which it is really present ;
although in examining the numerous Retepores in the collection I have throughout looked for
Loxosoma. My evidence goes to show that there is no restriction of a particular species of
Loxosoma to one species of Retepora; and the same species of "host" may bear more than
one species of Loxosoma. In the majority of cases the Entoproct is not the only commensal
organism, the Retepora being infested simultaneously with a Gymnoblastic Hydroid, with an
elongated hypostome bearing diffusely arranged tentacles. From the fact that this Hydroid
produces buds which appear to be destined to break off as free Medusae I refer it to the
genus Syncoryne. I have already called attention to the occurrence of the Loxosoma and
Syncoryne on Retepores in my Presidential Address to Section D at the Dublin meeting of
the British Association l). The Loxosoma is most commonly found on the frontal side of the
Retepore; but the specimens 337. J., which I have selected as the types of L. cirriferum,
occurred mostly on the basal sides or backs of the branches. I have usually, though not
always, found the Entoprocts on parts of the Retepores in which the polypides were in full
functional activity.
Most of the other species of Loxosoma appear to have an even number of tentacles.
In L. annulatum I have, however, counted nine with certainty in several specimens, and I have
been able to ascertain that the odd tentacle is situated distally. In one case there appear to
be 1 1 tentacles.
The stalk varies in length more than in most species. The extreme measurements which
I have made are 166 u. and 454 >j.. lts breadth is considerable, and may be as much as 51 >j..
The robust stalk and the rather narrow calyx give this species a very different appearance
from L. circulare, for instance.
In most species of Loxosoma the old bud is attached to the parent by the tip of its
stalk. In the present species the tip is prolonged beyond the point of attachment ; a feature
which was described by Claparède 3) and confirmed by Nitschk :') in L. kefersteinii. I have
had some difficulty in deciding how the foot-gland of the bud is related to this prolonged tip;
but the evidence of 247. E. seems to show that the attachment of the bud is at first quite
normal (fig. 3), the foot-gland running from the point of attachment, along the oral side of
the stalk, to the commencement of the calyx. At this region there is at first an indentation
between the stalk and the calyx ; but this soon disappears, and the opposite end of the stalk
begins to grow towards the lophophoral end of the parent. The attachment or "umbilicus" of
the bud then lies on that side of its stalk which is opposite to the surface containing the last
remains of its foot-gland (figs. 4, 5).
ij Harmer, S. F., 1909, Report Brit. Ass., Dublin Meeting, 1908, p. 721.
2) 1 ! .1 u ini. E., 1870, "Beitr. Anat. u. Entw. d. Seebryozoen", Zeitsch. wiss. Zool., XXI, p. 171, PI. X, tig. 4.
3) Nitsche, 11., 1875. "Bau u. Knospung v. Loxosoma ktl\i sicinii'\ Ibid., XXV, p. 454; and 1S75, "t'eb. d. Knospung d.
1 !.. XXV, Suppl. Bd, p. 37(1, PI. XXV, iig. 4.
It would appear, from this singular arrangement of the end of the young stalk, that
L. annulatum is related to L. kefersteinii\ but the nuniber of its tentacles is considerably
smaller than in that species, and buds are developed less profusely.
The following specimens probably belong to the present species ; but as neither of thcm
is associated with a Retepore I have thought it best to record them separately :
416. J. On Schizoporella, 416. G. (slide). Stat. 321. 6° 5'. 5 S., 1 i3°3o' E., 82 Metres; fine grey mud.
130. P. With Valkeria tuberosa, 130. G.2, Stat. 164. t°42'.5. S., I30°47'.5 E., 32 Metres; sand,
small stones and shells.
Each of the above specimens is represented by a single individual.
416. J. is attached to the frontal wall of a zooecium of a species of Schizoporella. It is
seen sideways, in a position imfavourable for examination. lts general shape and size correspond
well with the specimens from Retepores. lts stalk shows distinct indications of annulation, and
has an expanded disc of attachment, without foot-gland. There is an advanced bud, on one
side, the tentacles of which appear to be 9, or about 9, in number.
1 30. P. occurs on a Valkeria slide, but the Ctenostome had no doubt been growing
on some other object ; and its association with the Loxosoma on the same slide may have
been purely accidental. There is not sufficiënt reason to assume that it was really the "host"
of the Loxosoma. The specimen appears to agree closely with the preceding individual, and is
probably correctly referred to the present species.
4. Loxosoma velatum n. sp. (PI. I, figs 8 — 10).
24. D. On Retepora, 24. A. Stat. 53. Bay of Nangamessi, Sumba, o — 36 Metres; coral-sand ;
near the shore, mud.
Calyx (without the expanded membranous margins) about 205 u. long, and (with the
marginsj about 198 u. broad; expanded laterally into a very broad, thin, velate margin, which
extends on to the distal part of the stalk. Length of calyx plus velum, 320;/. In young
individuals (fig. 9), in which the velum is not yet differentiated, the stalk appears to expand
widely as it joins the calyx. The calyx bears a varying number (less numerous in young
specimens) of cirriform organs, some of which may be branched. 4 or 5 of these organs are
generally present on each side ; the distal pair forming horn-like structures in the region of
the lophophore, and the others being borne by the edges of the velum. Buds not more than
three at a time, a nearly mature bud being accompanied by a small bud on its own side and
a bud of intermediate age on the opposite side. Tentacles apparently about 14 — 16, the
lophophore transversely elongated in the retracted condition. Stomach with the lateral lobes
barely indicated. Sexes separate ; a pair of testes or a pair of ovaries being present. Stalk,
without its velate portion, longer than the part of the calyx which contains the lophophore and
stomach; with small projecting papillae laterally; ending basally in a small disc of attachment:
with no foot-gland in the adult, but a well developed gland in the buds.
Found attached to a species of Retepora.
13
The specimens here described were found with L. circulare, from which they are easily
distinguished by the velate expansion of the adult, or by the expanded junction of the stalk
and calyx, due to the development of the velum, in immature individuals (fig. 9). The velum
is not unlike the similar expansion in L. annelidicola Van Ben. and Hesse *), from which
species L. velatum differs conspicuously in other respects, as for instance in the general form
and in the presence of a foot-gland in the bud, a structure which, according to Prouho -), is
absent at all stages in L. annelidicola.
5. Loxosoma cirriferwm n. sp. (PI. I, figs 11 — 13).
Type. 337. J. On Retepora, 337. C. Stat. 71. Makassar, o — 32 Metres; mud, sand with mud, coral.
(And on the Retepora slide, 337. C.2).
193. E. On Retepora, 193. A.3 Stat. 274. 5°28'.2S., I34°53'.9E. 57 Metres; sand and shells,
stones.
193. G. On Retepora, 193. F.1. Stat. 274. Ibid.
297. D. On Retepora, 297. A.1. Stat. 273. OffPulu Jedan, Aru Islands, 13 Metres; sand and shells.
[The addition of a numeral to the symbol representing the Retepore indicates, as in
other cases, that the Loxosoma is be found on the slide so designated. Thus 193. E is to be
found on the Retepora slide marked 193. A.8].
Calyx relatively large, reaching a length of 2S0 — 310 u. , and a breath of 270 fx; broader
distally than proximally. lts margin bears a varying number of cirriform appendages, commonly
dilated proximally and with a filiform termination, sometimes branched. Tentacles 14 — 18 in
number, less numerous in young individuals. Stomach more or less globular, without distinct
lateral lobes. Sexes separate ; the females with a short row of eggs in the ovary, and carrying
as many as six or seven embryos in their vestibule (fig. 12). Buds as many as three at a
time ; an old bud being associated with a young bud on its own side and one of intermediate
age on the opposite side. The last attachment of the bud to the parent is by the tip of the
stalk, the foot-gland running along the oral side of the stalk as far as the commencement of
the calyx. Stalk variable in length (doubtless partly as the result of varying contraction) ;
sometimes slightly shorter than the calyx, but sometimes considerably longer, and reaching a
length of 400 [J. ; terminating proximally in a slightly marked disc of attachment , without
foot-gland.
The specific name which I have chosen for the present species indicates what I regard
as one of its most distinctive features. The apparent variability of the occurrence ot the cirri
may be due to the fact that these structures are deciduous or easily affected by reagents.
Nickerson 3) has given reasons for believing that the "flash organs" of L. davenporti are not
permanent structures, but are deciduous, while others may develop later ; and he comments
on the complete absence of these structures in some individuals. While not expressing himself
1) Cf. Prouho, H., 1S91, Arch. Zool. Exp., (2) IX, PI. V.
2) Ibid., p. 110.
3) Nickerson, W. S.. 1901, J. Morphol., XVII, pp. 357, 35S, PI. XXXII, fig. 11, PI. XXXIII, figs 16—19.
14
15
positively on the subject, he is inclined to regard them as being glandular in nature. The cirri
of L. cirriferum appear to be of the same nature as the "flash organs" ; but I think that
they are tactile organs, which are probably of special use in enabling the Loxosoma to avoid
injury by the formidable avicularia of the Retepore.
Cirriform organs have been described in certain other species of Loxosoma. The structures
shown at the margin of the calyx in L. karmeri by Schultz a) are probably of this nature.
More or less similar organs have been described by Assheton 2) in L. loxalimim and L.
saltans. The two "posterior sense-organs" of which I have given an account3) in L. crassi-
cauda Sal. are perhaps more specialised organs of the same general nature; while more or
less similar structures have been described by Vogt '*) in L. phascolosomatum, by Nnxs 5) in
L. brumpti and by Mortensen °) in L. antedonis. But these species are all sufhciently different
in other respects from L. cirriferum to make it impossible to refer the 'Siboga' specimens to
any of them.
Young individuals of L. cirriferum, whether advanced buds or recently detached free
forms, show no tracé of the cirri, which appear to increase in number as the individual grows
larger. Thus the small male specimen shown in hg. 1 3 has only two pairs of cirri. The small
specimens also differ from the large ones in another important particular, since they appear to
have a smaller number of tentacles. As in many other cases, in this genus, I have found it
difhcult or impossible to count the tentacles certainly. I have found evidence, however, that
the full number of the tentacles is 18, although other specimens, smaller in size, seem to have
no more than 16 or 14; or perhaps in some cases 12. It thus appears that, in the present
species at least, there is a progressive increase in the number of cirri and tentacles after the
individual has become free.
L. cirriferum differs from the species next to be described in the fact that the females,
or the older ones at least, carry a number of embryos simultaneously in their vestibule. Cor-
related with this fact is the occurrence of a short row of eggs (about 3) in the ovary (figs 11, 12);
other species frequently having only one recognisable egg at a time in each ovary. Male spe-
cimens are usually smaller than females, although I have found a male almost as large as the
largest females. I find no evidence of protandry ; and eggs may be produced by females which
are no larger than the male shown in fig. 13.
The buds in this species occupy a relatively proximal position. In the smaller indivi-
duals they are developed opposite what may be considered the middle of the stomach ; but
this organ has a considerable extension on the lophophoral side of the insertion of the buds,
which are thus situated nearer the proximal end than the distal end of the stomach (fig. 13).
In old individuals (fig. 1 2) the proximal insertion of the buds is very marked. It will be
obvious from the figures that buds are borne indifferently by males and females.
1) Schultz, E., 1895, Trudui St Peterb. Obschch., XXV, Sect. de Zool. & Physiol. Pt II, text-fïg. on p. 51.
2) Assheton, R., 1912, Quart. J. Mier. Sci., LVIII, pp. 118, 124, PI. VI, figs 1, 19.
3) Harmer, S. F., 1885, Quart. J. Mier. Sci., XXV, p. 273, PI. XIX, fig. 1.
4) Vogt, C, 1876, Arch. Zool. Exp. V, p. 312. PI. XI, fig. 3, PI. XII, fig. 1.
5) NlLUS, G., 1909, Trudui St Peterb. Obschch., XL, Livr. 1, N° 4, e. r. séances, p. 16S, text-fig. on p. 165.
6) Mortensen. Th., 191 i, "Danmark-Exsped. GrOnlands Nordostkyst", V, N° 8, p. 400, PI. XXVI, figs 2—4, 0.
IS
i6
6. Loxosoma circulare n. sp. (PI. I, figs 14 — 16).
Type. 24.15. On Retepora, 24. A. Stat. 53. Bay of Nangamessi, Sumba, o — 16 Metres; coral-sand;
near the shore, mud.
281. E. On Retepora, 281. D. Stat. 43. Off Pulu Sarassa, Postillon Islands, o — 36 Metres; coral.
? sp. 358. P). On Retepora, 358. A. Stat. 240. Banda anchorage, 9 — 45 Metres; black sand, coral,
Lithothamnion.
Calyx smaller than in the preceding species, usually from 150 to 190 a long and from
125 to 150 ij. broad ; nearly circular in the retracted state of the tentacles. Two pairs of tactile
appendages, at the margins of the proximal half of the calyx. Tentacles probably 12. Stomach
with indications of lateral lobes. Sexes separate ; the females with a single large yolky egg in
the ovary and with not more than two embryos in the vestibule (fig. 14). Buds as in L. cir-
riferum, provided with a foot-gland. Stalk usually slightly shorter than the calyx, from which
it is sharply marked off; from 1 10 to 155 u. in length in the preserved specimens, and 26 —
32 ij. broad; terminating in a well marked disc of attachment, without food-gland in the
adult state.
The specific name which I have chosen for this Loxosoma refers to the nearly circular
outline of the calyx, when the tentacles are retracted.
I was at first in considerable doubt whether this form should be regarded as distinct
from L. cirriferum. The difference in the female specimens of the two forms seems, however,
to justify its separation. Only one egg is produced at a time in each ovary ; and there are
accordingly only two embryos simultaneously in the vestibule. Both eggs and embryos are larger
than in L. cirriferum, as will be seen by comparison of the figures given of the two species.
The possibility is perhaps not excluded that the specimens here described are young forms
of L. cirriferum ; and that with increasing size and age the number of the eggs increases
and their size diminishes; but I do not think that this view is, a probable one.
The specimens 358. B. are distinctly larger than the ones of which the measurements
are indicated in the diagnosis, their calyx reaching a length of 288 \j. and a breadth of 224 u..
I am not sure that they are rightly referred to the present species.
7. Loxosoma pusillum n. sp. (PI. I, figs 19, 20).
Type. 133. G. On Retepora, 133. B. Stat. 164. i°42'.5 S., 130° 47.5 E. 32 Metres; sand, small stones
and shells.
133. C. On Retepora, 133. B. Stat. 164. Ibid.
193. H. On Retepora, 193. C.2, C.3. Stat. 274. 5°28'.2S., I34°53'.9E. 57 Metres; sand and
shells, stones.
108. AM. On Retepora, 108. B.2 Stat. 144. North of Salomakiëe (Damar) Island, 45 Metres;
coral bottom and Lithothamnion.
Size of individuals small, the calyx commonly not more than 100 [j. long and about
75 ij. broad; in larger specimens as much as 147 a long and 128 u. broad; resembling that of
L. circulare in general shape; and sharply marked off from the stalk. Cirriform organs minute,
one pair being present opposite the proximal half of the stomach. Tentacles 8 — 10, but some-
times not more than 6 in young specimens. The thickened epithelium of the stomach extends
16
17
round the sides of die organ, leaving only a small proximal part lined by thin epithelium. Sexes
separate ; the mature female showing a large yolky egg in one of its ovaries, and carrying one
embryo in the middle of its vestibule. Buds situated rather proximally, as many as two pairs
being present simultaneously; a foot-gland developed in this stage. Stalk varying in length
according to its degree of contraction; when stretched, as much as 16011 long and ig u broad;
vvhen contracted, not more than i 10 a long and accordingly broader (32 u,). The stalk has a
well marked disc of attachment, without foot-gland in the adult state.
This form has a considerable resemblance to L. circitlare, of which it may indeed be
a dwarf variety. The number of tentacles is, however, smaller than in that species; and only
one embryo appears to be developed at a time. I have, moreover, only found one pair of
cirriform organs. These differences are, perhaps, not inconsistent with the view that it is a
small form of L. ciradare ; but it seems to me preferable to describe it as a distinct species.
In size and number of tentacles the present species resembles L. shiiteri, which differs
from it in its narrower calyx. It is, besides, very improbable that the same species of Loxosoma
would occur on two "hosts" so different as a Sipunculid and a Retepore.
Loxosoma spp. incert.
89. B. On Retepora, 89. A. Stat. 116. West of Kwandang Bay, entrance, 72 Rletres; fine sand
with mud.
150. F. On Retepora, 150. A. Stai. 204. Between Islands of Wowoni and Buton, 75 — 94 Metres;
sand with dead shells.
108. AG. On Retepora, 108. B. Stat. 144. North of Salomakiëe (Damar) Island, 45 Metres;
coral-bottom and Lithothamnion.
216. C. On Retepora, 216. A. Stat. 285. South coast of Timor, 34 Metres; Lithothamnion.
231. B. On Retepora, 231. A. Stat. 305. Solor Strait, off Kampong Menanga,_ 113 Metres;
stony bottom.
410. E. On Retepora, 410. B. Stat. 305. Ibid.
On Retepora. Singapore, R. HAXITSCH Collection, Mus. Zool. Cambridge, Reg. Apr. 10, 1900.
Specimens of Loxosoma were observed on the Retepora indicated above ; but as
microscopic preparations were not made it would be unsafe to attempt to refer them to their
respective species.
8. Loxosoma troglodytes1) n. sp. (PI. I, figs 17, 18).
Type. 131 T., in compensation-sac of 131. A., Lepralia celleporoides. Stat. 164. i°42'.5 S., I30°47'.5 E.,
32 Metres; sand, small stones and shells. (Also on mounted slides of 131. A.).
142. D., in the same position and from the same Station (on slide 142. A., L. celleporoides).
322. 1., in the same position, in 322. F., L. celleporoides. Stat. 50. Bay of Badjo, W. coast of
Flores, o — 40 Metres; "mud, sand and shells, according to locality".
Size small. Form of the retracted animal pear-shaped, the narrower part being repre-
sented by the stalk, which passes almost continuously into the calyx. Total length, including
the stalk, about 185 — 192 fx; breadth of the calyx 102 — 109 u. The calyx bears two or more
1) TfwyAoJüT^i;, "one who ci'eeps into holes, a cave-dweller".
17
S1BOGA-EXPEDITIE XXVUI0.
i8
tactile processes at its distal end. Retracted lophophore nearly circular, with 9 tentacles. The
contracted stalk is about the same length as the calyx, its wal] possessing a layer of strong
longitudinal muscles : proximal end of stalk more or less pointed, without foot-gland in the
adult state. Buds not numerous, not more than two present on the side hearing the oldest
bud. Gonads?
The Cheilostome in which this species of Loxosoma has been found is the form which
was described by Busk l) as Lepralia cclleporoides. The middle of the frontal surface of this
species is occupied by a very large "median pore", of variable shape. The pore, which is
shown by Busk in PI. XVII, fig. \b, is morphologically an "ascopore" 2), since I find that it
is the external opening of a large compensation-sac, which contains the Loxosoma. The occur-
rence of Entoprocta as normal inhabitants of the compensation-sac of a Cheilostome has not
previously been recorded, so far as I am aware. The unusually large size of the ascopore in
L. cclleporoides renders the compensation-sac an eligible dwelling-place for the Loxosoma, which
has no difficulty in protruding its calyx through the one of the lobes of the ascopore to the
outer water, as shown by several individuals which have died in this extended attitude. It
need hardly be pointed out that the stalk is correspondingly elongated in such specimens.
In the retracted condition in which the great majority of the individuals have been
preserved, the proximal end of the stalk is usually attached to one side of the compensation-
sac, the calyx lying just underneath the ascopore, through which its tactile processes are com-
monly visible. The aboral surface of the Loxosoma lies uppermost, so that the orifice of the
vestibule faces the cavity of the compensation-sac. In many parts of the host the majority of
the zooecia, or all of them, contain Loxosoma ; from one to three individuals occuring in a
single compensation-sac, the calyces all radiating in the direction of the ascopore.
The Loxosoma, whose cave-dwelling habit I have indicated by the specific name sug-
gested, is not the only organism which has taken advantage of the large size of the ascopore ;
since, in many zooecia, one or more of the tubes of the Infusorian Folliculina are attached
to the wall of the compensation-sac, near its orifice ; the tube projecting through the ascopore
to the outer water. Where Folliculina occurs I have not found the Loxosoma, although some
zooecia contain neither of the two organisms. The Loxosoma is usually absent in zooecia which
have lost their polypides.
In many cases the tactile processes of the Loxosoma are two in number, disposed
symmetrically, as in figs 17 and 18. But in other cases three may be present; or there may
be two tactile processes, both situated on one side of the middle line ; or four, symmetrically
disposed.
In many species of Loxosoma the tentacles are present in an even number. But in the
present species nine tentacles were counted in all cases where the number was definitely
ascertainable. Of these, four tentacles belong to the proximal end of the lophophore, and the
odd tentacle is the distal one.
The form of an old bud is shown in fig. 17, where a young bud is seen on the distal
ij Bi -k. ('.., 1884, 'Challenger' Report, Pt XXX, p. 142.
2) LEVINSEN, G. M. R., 1909, "Morph. and Syst. Stud. Cheilostomatous Biyozoa", Copenhagen, p. V.
18
19
side of the base of the older one. I have been unable to ascertain whether a foot-gland is
present in the bud or not ; and I cannot give any reliable information with regard to the
gonads.
9. Loxosoma breve n. sp. (PI. I, figs 29 — 31).
? Loxosoma singulare Waters, 1914, "Mar. Fauna Brit. E. Afr." Proc. Zool. Soa, p. 855 (on
Sckizoporella nivea, mostly 011 the opercula).
131. U. On Sclücoporclla, 131. B.3 (Type), 131. B.1; and on Cellepora, 131.Q.2, Q.3. Stat. 164.
i°42'.5 S.j i30°47'.5 E. 32 Metres; sand, small stones and shells.
Size small, total length of the preserved and retracted specimens not exceeding 2 30 ;;..
Of this, the largest specimen found, the calyx m'easures 147 p, in length and the stalk S 3 y. :
the calyx is 128 u. broad, and the stalk is as much as 70;;. broad. In other specimens which
are better extended the stalk is more elongated, and thinner, measuring 134,'J. in length and
only 16 u in breadth. The calyx varies in form according to its degree of contraction, but is
oval in those which appear to be best preserved. Several short tactile processes occur on the
distal part of the calyx. Number of tentacles 9. Stomach globular. Buds produced in small
numbers (not more than one at a time found). Sexes apparently separated, a single individual
showing a pair of ovaries, each with a fairly large egg. Stalk of adult terminating in an
expanded disc of attachment, without foot-gland.
I have found this species on slides of a Sckizoporella l), and a Cellepora, both from
the same Station ; several individuals being present on each slide. In the case of the Sckizo-
porella^ the attachment is to the outer side of the operculum of the "host" ; and the specimens
show distinct tactile processes. The Cellepora preparations had been decalcified ; and it is
uncertain how the Loxosoma were attached. These specimens show no tactile processes; but it
does not seem necessary to separate them from the others on that account.
The species appears to be related to L. froglodytes, but is distinguished from it by
the want of the conspicuously strong longitudinal muscles of the stalk which are characteristic
of that form.
10. Loxosoma subsessile n. sp. (PI. I, figs 32, 33).
260. G. On Conescharellina, 260. D.1. Stat. 318. 6°36'.sS., H4°S5'.5E. 88 Metres; fine yellowish
grey mud.
Size minute, total length not exceeding about 128 <j. in the preserved material. Calyx
not much broader distally than proximally, the well marked lateral lobes of the stomach being
accommodated by a corresponding breadth of this part of the calyx, which measures from 90
to 1 10 y. in length and 74 to 94 <j. in breadth. Tentacles 8. Buds one on each side. Stalk
extremely short, about 26 y. long; much narrower than the calyx, from which it is sharply
separated ; without foot-gland or marked disc of attachment.
1) L. singulare has been recorded from a Sckizoporella by G. A. CORNISH, 1907, "Rep. Mar. Tol. Canso. N. S. [Xova Scotia]",
39th Ann. Rep. Dep. of Marine and Fisheries, Fisheries Branch (Ottawa), p. 80.
19
20
The specific name refers to the shortness of the stalk. I have found this species on
one ocasion only, on a species of Conescharellina. lts minute size and general proportions of
calyx and stalk distinguish it satisfactorily from the other 'Siboga' species. I am able to give
the number of tentacles, with some confidence, as 8. In this respect the present species agrees
with L. nitschei Vigelius1); which it also resembles (judging by the figures given by that
author) in general form. The "height" is given by Vigelius as about 150a; but the original
figures give no indication of the strongly developed lateral lobes of the stomach which seem
to be characteristic of the 'Siboga' species. L. nitschei was found by Vigelius on Menipea
ternata ; the locality being Barents Sea, as may be inferred from the localities of the "host"
given in the same paper.
I have recently had the opportunity of examining specimens of Loxosoma found by Miss
R. E. Roper in great abundance in a tank in the Dove Marine Laboratory at Cullercoats,
Northumberland, and recorded by her 2) as L. nitschei. If this determination be correct, as I
am disposed to think, the 'Siboga' species is probably distinct from the form described by
Vigelius: --a conclusion which has much a priori probability from the widely distant localities
of the two forms under consideration. Miss Roper has commented on the fact that the total
length recorded by Vigelius is less than half that of the living Cullercoats material ; and in this
respect I can confirm her observations. It may be worth while to record my own experience
of this species, as bearing on the value to be ascribed to measurements of specimens of
Loxosoma which have not been preserved with special care. The specimens of Miss Roper's
species examined by me were received alive, and were narcotised with cocaine, in order to
preserve them with their tentacles extended. Even in the narcotised condition there is some
clifficulty in killing them without contraction. In some of the individuals in which contraction
had occurred, the general proportions were very similar to those figured by Vigelius, the stalk
being much shorter than the calyx. It must be admitted that even in these cases the measure-
ments considerably exceed that given by Vigelius, the total length being not less than 371 ju,
in adult individuals; of which 243 u belong to the calyx and 128 a to the stalk. In others
which had been killed more suddenly with strong osmic acid the stalk measured as much as
320;;. in length, its breadth being noticeably less than that of the more contracted specimens.
The total length in an expanded specimen may be as much as 590 u.. It is thus obvious
that the stalk of a Loxosoma which has been preserved without being narcotised is likely to
be considerably shorter than the stalk of the living, fully extended animal ; although the
measurements of ordinary, preserved material may be of some value comparatively, since the
contraction probably takes place more or less equally in the majority of the specimens.
Although the Cullercoats specimens are thus considerably larger than those from Ba-
rents Sea described by Vigelius I think it is on the whole probable that they belong to the
same species. If this conclusion is justified, the 'Siboga' specimens here described differ from
1) VlGELtUS, W. J., 1SS2, "Cat. Polyzoa 'Willem Barents'", Nied. Arch. Zool., Suppl. Bd I, p. 19, Plate, figs 4(7, 4/S, 5. The
species is also recorded, doubtfully, by K. A. Andersson, on Flustra mtmbranaceo-truncata Smitt, from E. Greenland (Zool. Jahrb. Syst.
XVI, 1902, p. 555).
2) ROPER, R. E. "The Marine Polyzoa of Northumberland", Dove Marine Laboratory, Cullercoats, Report for the Vear ending
June 13'!', 1913, p. 56.
20
2 I
L. nitschei so greatly in size that they can hardly be regarded as belonging to the same
species. It must, however, be pointed out that the Cullercoats specimens agree with them in
the transverse elongation of the stomach : — a fact which would not have been suspected
from an inspection of Vigelius' rather diagrammatic figures of the Barents Sea Loxosoma.
11. Loxosoma loricatum n. sp. (PI. I, figs 21 — 25).
Type. 177. N., on Canda, 177. A. Stat. 257. Du-roa Strait, Kei Islands, o — 52 Metres; coral.
108. AJ., on Canda, 108. K. Stat. 144. Anchorage N. of Salomakiëe (Damar) Island, 45 Metres ;
coral bottom and Lithothamnion.
::279. G., on Canda, 279. B. Stat. 250. Anchorage orï Kilsuin, W. coast of Kur Island, 20 — 45
Metres; coral and Lithothamnion.
'204. G., on Canda, 204. A., Stat. 282. Anchorage between Nusa Besi and the N. E. point of
Timor, 27 — 54 Metres; sand, coral and Lithothamnion.
*270. M., on Canda, 270. E. Stat. 303. Haingsisi, Samau Island, o — 36 Metres; Lithothamnion.
224. B., on Canda, 224. A. Stat. 305. Solor Strait, off Kampong Menanga, 113 Metres; stony
bottom.
Size small, the total length not exceeding about 160 — 210 u.. The calyx, which is about
80 — 108 ij. broad, is covered on its posterior or aboral side by a well marked chitinous lorica,
about 1 30 — 1 60 \j. in length, and thus reaching nearly to the proximal end of the stalk. The
lorica is specially gibbons or even subcarinate in its proximal half, the distal half being more
flattened. It is sometimes marked by lines, longitudinal in the proximal half and concentric
with the distal margin in the distal half (fig. 25); but these may have been artificially pro-
duced by contraction. The lorica is almost conterminous with the calyx, since the stomach
extends nearly to its proximal end. The calyx bears a pair of minute projecting papillae,
probably tactile organs, at the sides of the lophophoral region (fig. 21); they are situated on
its anterior side, close to the edges of the lorica. Lophophore small, with 8 — 10 tentacles.
Stomach without lateral lobes, rather elongated. Buds few, not more than one (on one side
only) visible. Stalk extremely short, slightly overlapped by the lorica, and much shorter than
that structure ; ending proximally in a distinct disc of attachment, about 52^. broad, without
foot-gland. Gonads?
The specimens here described were all found on the same species of Canda (to be
described in a later part of this Report). Their position appears to be nearly constant, the
Loxosoma usually occurring on the frontal side of the "host" attached to the distal end of a
zooecium, near its outer margin (fig. 25). More rarely a Loxosoma was found on the back of
a branch. The vibracular seta of the host can be moved to the frontal side of the branch,
where it takes up a more or less transverse position, just distally to the point of attachment
of the Loxosoma. The stalk of the Entoproct often lies over the seta, the calyx being directed
towards the distal end of the Canda, and having its anterior or vestibular side facing the
branch. It may be presumed that the L.oxosoma is not seriously incommoded by the move-
ments of the seta ; but it is obvious that as it commonly lies transversely across it, it must
* The specimens marked with an asterisk were «.amined only with a low power, in spirit.
21
move into an upright position every time the seta is moved to the back of the branch. From
the relative positions of the Loxosoma and the seta, it might be supposed that the Entoproct
would sometimes be crushed between the seta and the frontal surface of the branch, when the
seta is brought round to the front. But I find no evidence that this actually happens, since
the seta always lies on the branch, when in the position referred to, with the Loxosoma
outside it. It may be supposed that the Loxosoma reacts very quickly to the stimulus of the
movement of the seta, or that it habitually holds itself in an erect attitude, so that it avoids
the danger of being injured to which it might otherwise be exposed.
In some parts of the Cauda, every zooecium has a single Loxosoma, in the position
above described. I have not found more than one Loxosoma on any zooecium.
The number of the tentacles seems to be usually 8, as in L. nitschei and Loxocalyx
cocJilear; but I think I am right in saying that it may sometimes rise to 10. I am unable to
decide whether a food-gland is present in the bud or not; and I have no reliable observations
about the gonads.
The present species is quite distinct from any other with which I am acquainted. In its
loricate condition it approaches L. cocciforme, described below ; but the lorica is less developed
than in that species, and does not extend far enough to cover the stalk completely v).
It may be remarked that the dried and shrivelled remains of the Loxosoma are some-
times quite recognisable in dry preparations of the Canda.
12. Loxosoma cocciforme n. sp. PI. I, figs 26 — 28).
112. L. On 112. D., Siphonicytara (Cheilostomata). Stat. 156. o°2o/.2S., i30°s'.3E., coarse sand
and broken shells. (Mounted on thê slides 112.D.).
Form of the calyx resembling that of a Scale-Insect, nearly circular in outline, but
becoming narrower at the proximal end, attached by a concave or flattened surface to the
outside of its "host", its opposite surface convex. Length of calyx about 220 — 270 *j., breadth
190 — 220 'x. Margin very thin all round. The calyx is covered, on its free surface, by a
thick cuticle. Stomach small, without distinct lateral lobes. Tentacles usually 10, but in the
larger individuals 12. Lophophore transversely elongated, the opening of the vestibule trans-
versely oval during retraction and situated on the attached or concave side of the animal.
Buds not numerous. Sexes apparently separate. Stalk extremely short, not nearly so long as
the calyx, by which it is completely concealed in the retracted state, in which condition the
long axis of the stalk is more or less at right angles to the attached surface. The stalk
terminates in a disc of attachment, without foot-gland.
This peculiar species has the form which is sufficiently indicated by the specific name
which I propose for it. It was found closely adherent to the surface of the Cheilostome on
which it was growing, in most cases just on the distal side of a peristome. The majority of
1) A chitinous aboral shield on the calyx has been described by Annandale (1908, Fauna ISrackish Fonds Port Canning",
Pt VII, Ree. Ind. Mus., II, Pt I, p. 14) in the species described by him as Loxosomatoiiles colonialis; which is, however, quite distinct
from the 'Siboga' species in other respects.
22
23
the individuals were on the "back" of the host; but some were found on its "front" surface.
The form of the body may be regarded as an exaggeration of that of L. annelidicola Van
Beneden and Hesse ; as represented by Prouho l). But while in L. annelidicola the marginal
expansion is incomplete at the proximal end, leaving an emargination through which the short
stalk projects, the proximal end of L. cocciforme is not emarginate and completely covers the
retracted stalk. The two species appear to be related to one another, although I have not been
able to ascertain whether the bud of L. cocciforme possesses a foot-gland, a structure which
is completely absent, even in the buds, according to Prouho a), in L. annelidicola.
The buds of L. cocciforme are obviously not numerous, as is indicated by fig. 26, in
which a single . moderately large bud is seen on one side, projecting beyound the margin of
the calyx of the parent-individual. Ovaries and testes appear to occur in different individuals; -
as I have found to be the case in all the other 'Siboga' species in which I have been able to
obtain definite evidence on the subject. In the specimen represented in fig. 28 several embryos
are seen inside the vestibule.
Waters3) has recorded L. singulare, Franz-Josef Land, from a depth of 234 fathoms ;
and states that this appears to be the greatest depth at which the genus has been found. The
present species, from 469 Metres (= about 256 fathoms), was obtained in even deeper water
than the record indicated.
Fam. 2. Pedicellinidae Johnston *).
There has been considerable difference of opinion with regard to the number of genera
which should be recognised in this Family. While the type-genus possesses a stalk which is
uniformly muscular and has no differentiated enlargement at its base, a number of other species
are distinguished by having a specially contractile portion at the origin of the stalk from the
stolon. Several of the 'Siboga' species are thus characterised ; but the following species comes
within the original genus.
Pedicellina M. Sars.
1835, "Beskr. og Iagtt. over Bergenske Kyst levende Dyr", p. 4.
This genus is distinguished from Barentsia by having a stalk of uniform thickness, not
differentiated proximally into a special muscular enlargement.
1) Prouho, H., 1891, Arch. Zool. Exp. (2) IX, PI. V, figs 3, 7.
2) t. cit., pp. 110, 113.
3) Waters, A. W., 1904, "Bry. Franz-Josef Land", II. J. Linn. Soc. Zool., XXIX, p. 181.
4) In the 'Challenger' Report, Part 50, 1886, p. 40, Busk gives "Pedicellinidae Hincks"; referring apparently to "British Marine
Polyzoa, 1880, p. 563, where the Family is characterised. But, as shown by Busk's citations, Johnston, in his "History of the Britlsh
Zoophytes", Ed. 2, 1847, p. 381, had already used "Family Pedicellinae", referring to Gervais, 1837, as giving "Polypiaria Pedicellinea".
In Gray's "List of the Specimens of British Animals in the British Museum", Part I, 1848, p. 93, "Fam. Pedicellinidae'' is delinitely
used, and it is detined on p. 145. The facts thus seem to be that Johnston established a Family for the genus which M. Sars had
described in 1835, although Gray was the first to use it with the terminalion -idae, and to give it a diagnosis.
23
24
i. Pedicellina compacta n. sp. (PI. II, figs 13, 14).
293. B. Stat. 273. Anchorage off Pulu Jedan, Aru Islands, 13 Metres; sand and shells (on
293. A., Adeonelld).
Total length about 660 u., the stalk not much longer than the calyx. Tentacles about
12. Stalk relatively thick, uniform in diameter or diminishing towards the calyx; without spines.
Stolon-branches given off freely from the bases of the stalks, one or two usually present, on
each side, in this position. Sexes apparently separate.
The specific name refers to the comparatively thick and short stalk; though the con-
dition shown in figs 13 and 14 is no doubt that of contracted specimens. A diminution in
size of the stalk at its distal end, often observable, doubtless indicates that this part is less
contracted than the more proximal portion.
I have found this species only once, on a species of Adeonella, which in some parts
is profusely covered with it. The general character of the stolon is identical with that of other
species, a barren part alternatiug with a stalk-bearing portion. A chitinous diaphragm passes
across the stolon at no great distance from the base of a stalk ; and this is equally true of the
main stolon and of its lateral branches. The crowded arrangement of the individuals of the
colony is due partly to the fact that the barren stolon-segments are short, and partly to the fact
that lateral stolon-branches are given off very freely, as shown in fig. 14, where two are present
on each side. In individuals in which I have been able to count the tentacles, the number has
been 1 2 ; and I regard this as the typical number. In some cases, as in the specimen show in
fig. 13, the rectum has an erect position. I think that this condition is associated with the
development of a brood-pouch from the floor of the vestibule, since in young calyces in which
there is no brood-pouch the rectum is recumbent on the wall of the stomach. The gonads seem
to be female in all the individuals in which they can be made out; and this appears to indicate
that the sexes are separate and that all the specimens mounted belong to a female colony.
The measurements (in p.) of the individual represented in fig. 13 are: — length of
calyx, 300 ; of stalk, 360 ; total length, 660 ; breadth of calyx, 230; of stalk, 60 — 70.
The majority of the known species of Pedicellinidae belong to the genus Barentsia ;
and there are comparatively few which fall within Pedicellina s. str. Of these, the form which
most nearly resembles the present species in general appearance is P. australis, described by
Ridley l) from Sandy Point, Straits of Magellan, 7 — 10 fathoms. With this species it agrees
in the number of tentacles ("about 12"), but differs from it in size; the measurements given
by Ridley for his species including the following : — total length 2.5 mm., calyx 1 mm.
I have verified these measurements from the type slide (79. 12. 27.36) in the British Museum,
although I find that the total length, 2.5 mm., is obtained by including the breadth of the
stolon-segment from which the stalk arises. It thus appears that the calyx alone of Ridley's
species is longer than the total length of the 'Siboga' form: and the difference in size is so
marked that it seems impossible to refer the 'Siboga' species to P. australis.
1) Ridley, S. O., 1881, "Zool. Coll. 'Alert'". I'roc. Zool. Soc, p. 60. Pedicellina australis Jullien, is a Barentsia\ see the
account of B. discreta in the present Report, p. 29.
24
= 5
Another species which may be noticed in the present connexion is P. breusingi,
described by Studer x) from Kerguelen ld, zone of Florideae. This species is said to differ from
P. australis Ridley, in the number of its tentacles (16 — 18), in the irregular form of its stolon,
in the absence of diaphragms in the stolon, and in the shape of the calyces. It may be sur-
mised that the supposed absence of diaphragms was due to a mistake in the observation,
since it is not in agreement with what is known of other Pedicellinidae.
Barentsia Hincks.
Barentsia Hincks, 1S80, "Hydr. and Pol. Barents Sea", Ann. Mag. Nat. Hist. (5) VI, p. 285.
Ascopodaria Busk, 1886, 'Challenger' Rep., Pt. 50, p. 41.
Artliropodaria Ehlers, 1890, "Z. Kenntniss d. Pedicellineen", Abh. Göttingen, XXXVI, p. 144.
Gonypodaria Ehlers, 1890, t. cit., p. 144.
The Genus Barentsia, characterized by having a muscular enlargement at the proximal
end of its stalk, was established by Hixcks in 1SS0 for a new species, B. bulbosa ~), from
Barents Sea. This species differs from most of the forms which have subsequently been added
to the genus in the fact that the stalk gives rise laterally to new stalks-, a feature which is
also shown by B. variarticulaia Andersson s) and by Gonypodaria ramosa Robertson 4). In
1SS4 Hincks5) instituted a new genus, Pedicellinopsis; but he subsequently") expressed the
opinion that it was synonymous with Barentsia. In 18S6 Busk defined the genus Aseopodaria,
admiting its identity with Barentsia, but justifying his use of the name partly on the ground
that he had used it as a IMS. name in 1879, and that it had been mentioned in that year
by Allmax. Allmax's reference to the genus is, however, unaccompanied by any diagnosis,
so that Ascopodaria was a nomen nudiun when HlNCKS described Barentsia. In 1890 Ehlers
introduced the two genera Artliropodaria and Gonypodaria, the former characterised by its
jointed stalk.
The genera thus described have been criticised by various authors. Ritchie '), in a
recent paper, has given reasons for regarding Artliropodaria and Gonypodaria s) as synonyms
of Barentsia, on the ground of the variation, in the characters which have been used to
distinguish those genera, observable within the limits of a single species ; and he suggests that
any species of Barentsia can probably give rise to a joint in its stalk. This is perhaps going
bevond the evidence, since jointing has not been described in by any means all the species.
But that the stalk can become jointed in certain cases is clearly shown by many
observations, including his own here referred to. In two species the faculty of forming joints
1) Studer, T., 1S89, "Forschungsreise •Gazelle'", III. Theil, Zool. i; Geol., p. 140.
2) See also the descripticn,1 by \V. J. VIGELIUS, 1S84. of the same species in "Die Bryozoen 'Willem Barents1'', Bijdr. tot de
Dierkunde, XI, p. 85, figs on PI. VIII.
3) Andersson, K. A., 1902, "Bry. schwed. Exp.", Zool. Jahrb. Syst. XVI, p. 557. It seems doubtful whether this species is
really distinct from B. bttlbosa.
4) Robertson, A., 1900, "Stud. Pacific Coast Entoprocta", Proc. Cal. Acad. Sci. (3) Zool. II, X° 4, p. 337.
5) Hincks, T.. 1SS4. "Contr. Gen. Hist. Mar. Pol.", XIII, Ann. Mag. Xat. Hist. (5) XIII, p. 364.
6) Ibid., 1893, Appendix, Ann. Mag. Xat. Hist. (6) XII, p. 140.
7) Ritchie, J., 1911, "On an Entoproctan Polyzoon . .", Trans. R. Soc. Edinburgh, XLVII, Pt 4, p. 840.
8) Waters (1904, "Rés. Voy. Belgica", "l'.rv.", p. 100) had previously expressed this opinion with regard to Gonypodaria.
25
SIBOGA-EXPEDITIE XXVIII a. 4
2 6
leads to a specially regular transverse segmentation of the stalk. These are (I) B. benedeni
Foettinger; which is discussed by Ritchie ; and (II) Vrnatella gracilis Leidy. In the latter case
in particular it is clearly shown from the descriptions of Leidy l) and Dayexport -) that the
segmented condition is the result of constrictions formed in a stalk which is at first like that
of Pedicellina. In both B. benedeni, which constituted the type of Ehlers' genus Arthropodaria,
and l 'matcl/a, the stem-joints may give rise to buds which become new individuals, in much
the same way as in />'. bnlóosa, the genotype of Barenfsia. It is not impossible that the con-
dition seen in Pedicellinopsis is another development of the same process, if it be permissible
to regard the erect sterns which bear the Barentsia-Wke. individuals as modihed stalks of indi-
vidual units of the colony. Of this there is no direct evidence, so far as I am aware ; but the
account given by Busk 3) of the structure of the main "sterns" (see particularly figs 10 and 11
of his PI. IX) suggests that this may be their real meaning. P. fruticosa would, on this view,
be strictly comparable with B. bulbosa, but distinguished from it by the much more profuse
development of lateral buds from an original stem.
Urnatclla, which is referred by Davenport ') to the Pedicellinidae, doubtless deserves
generic recognition, from the peculiar characters of its stolon, represented by a basal plate of
attachment ; as well as by reason of structural characters of the calyx. The question whether
Pedicellinopsis should be regarded as a valid genus appears to me a more open one ; since
the general architecture of the colony is very different from that of the forms usually referred
to Barentsia. I have accordingly not included it in the synonymy at the head of this account.
But whether P. frziticosa should be included or not in Barenfsia, this genus includes a con-
siderable range of modifications of the stalk, as follows :
(I) stalk simple, without joints or lateral buds (usual form of B. gracilis M. Sars) ;
(II) stalk unjointed, but producing lateral buds (B. bnlbosa Hincks; B. variarticitlata Andersson);
(III) stalk jointed, bearing lateral buds (Gojiypodaria raniosa Robertson) ;
(IV) stalk more regularly jointed, with few or no lateral buds {Arthropodaria benedeni Foettinger).
These are probably to be understood merely as different conditions which ma}' be
assumed within the genus, and not as subgeneric groups ; since, as Foettinger r') and Ritchie
have shown, the stalk of B. gracilis (assuming, with Ritchie, that this species includes B.
belgica Van Ben.) may be unsegmented or segmented; and since the difterences between my
third and fourth groups are mainly difterences of degree.
It may not be out of place to enter a protest in this place at the use of the word
"polypide", by certain authors, to signify the calyx of a member of the Pedicellinidae. The
calyx consists of the body-wall of this part of the animal, including the alimentary canal and
nther organs. The term "Polypide" was introduced by Allmax c), at the Meeting of the British
i) Leidy, J., 1884, ^Urnatella gracilis"^ J. Acad. Nat. Sci. Philadelphia, (2) IX, p. 9.
2) DAVENPORT, C. B., 1893, "On Urnatella gracilis", Buil. Mus. Comp. Zool. Harvard, XXIV, p. 19.
3) Husk, G., 1886, p. 42.
4) Davenport, C. B., t. cit., p, 30.
5) Foettinger, A., 1886, "Sur 1'Anat. des Pédicellines de la cute d'Ostende", Arch. de Biol., VII, p. 301.
6) Ali.man. G. J., 1851, Rep. 20"' Meeling Brit. Ass., p. 307; see also his uMono^r. Fresh-Water Pol.'', 1S56, pp. 8, 41, 42.
26
27
Association held at Edinburgh in 1850, for "the retractile portion of the Polyzoa"; or, in
other words, for the alimentary canal, the tentacles with their sheath, and the ganglion, of the
Ectoprocta. The term "zooecium", in the sense in which it is now used, was employed later (as
"zoocecium") by Smitt '). Although the idea, once current, that the polypide and the zooecium
represent two different kinds of individuals has long been given up, the terms are in almost
universal use in describing Polyzoa. Nitsche 2) indeed expressed the opinion that the calyx of
an Entoproct represented the polypide only of Ectoprocta; but it is more in accordance with the
present state of our knowledge to regard a Pedice//iua-'md'w\dua.\ as comparable with an entire
unit (zooecium -f polypide) of one of the Ectoprocta. The term "polypide" if applied to the
Entoprocta should thus be used for the alimentary canal, tentacles and certain other organs which
it might be difhcult to particularise; the body-wall of the calyx, stalk and the corresponding
stolon-segment being the morphological equivalent of the zooecium of Ectoprocta.
1. Barentsia gracilis M. Sars3). (PI. II, fig. 12).
Pedicelllna gracilis M. Sars, 1S35, "Beskr. og Iagtt." (Bergen), p. 6, PI. I, lïgs 2a, 2b (Bergen).
Pedicellina gracilis Smitt, 1S71, "Krik förteckn.", V, Öfv. K. Vet.-Ak. Förh." XXVIII, p. 1133
(Sweden, Spitsbergen; nodes of stalk mentioned).
Pedicellina gracilis Hincks, 1880, "Hist. Brit. Mar. Pol.*', p. 570, PI. LXXXI, figs 4 — 6.
Pedicellina gracilis Hincks, 1884, "Pol. Q. Charlotte Islands", Ann. Mag. Nat. Hist. (5) XIII,
p. 208 (Queen Charlotte Islands).
Pedicellina gracilis Levinsen, 1894, "Zool. Danica", IV, 1 Afd., "Mosdyr", p. 96, PI. IX, fig. 30
(Denmark ; nodes of stalk figured).
Pedicellina gracilis Hilgendorf, 1898, "Occ. Pedicellina N. Zealand", Trans. N. Zeal. Inst.,
XXX (N. S. XIII), p. 218, PI. XXII, fig. i (New Zealand; nodes of stalk figured).
Barentsia gracilis Hincks, 1887, "Pol. Adriatic", Ann. Mag. Nat. Hist. (5) XIX, p. 312 (Adriatic;
nodes of stalk mentioned).
Barentsia gracilis Hincks, 1889, "Pol. St Lavvrence", Ann. Mag. Nat. Hist., (6) III, p. 432,
PI. XXI, figs 10, lOrt (St Lawrence).
Barentsia gracilis Carus, 1889, "Prodr. Zool. Med.", II, p. 53.
Barentsia gracilis Kirkpatrick, 1890, "Hydr. Pol. China Sea", Ann. Mag. Nat. Hist. (6), V,
p. 17 (Tizard Bank, betvveen Philippine Islands and Malay Peninsula, 27 fathoms).
Barentsia gracilis Ostrooumofï, 1896, "Otchet Exp. 'Selianik' ", Buil. Ac. Imp. St Pétersb.,
(5) V, p. 58 (Sea of Marmara).
Barentsia gracilis Calvet, 1902, "Bry. Cettc", Trav. Inst. Zool. Montpellier, (2) Mem. 11,
p. 94 (Cette).
Barentsia gracilis Waters, 1904, "Rés. Voy. Belgica", Zool., "Bryozoa", p. 100 (Queensland).
Barentsia gracilis Waters, 1910, "'Rep. Mar. Biol. Sudanese Red Sea", J. Linn. Soc, Zool.,
XXXI, p. 251 (Red Sea).
Barentsia gracilis Ritchie, 191 1, "On an Ent. Pol.", Trans. Roy. Soc. Edinb., XLVII, Pt 4,
pp. S41, 845, Plate, figs V — VII (Ostende, Isle of Man; nodes of stalk figured).
1) SMITT, F. A., 1865, "Krit. förteckn." I, Öfv. K. Vet.-Akad. Förhandl., X" 2, p. 115, note; in place of "djurhus" of a
previous paper. It is not generally remembered that it had been used, many years earlier, for the animals themselves, by LAMOUROUX, whose
actual words are "Les Polypiers, qui seraient mieux nommés peut-être Zoacies ou Synzoaciphytes" ("Hist. Pol. Cor. Flex.", 1816, p. xli).
2) Nitsche, H., 1871, "Beitr. Kenntn. Bry.", 111, Zeitschr. wiss. Zool., XXI, p. 483.
3) Here, as in other cases, references to the synonymy will be found in Miss E. C. Jei.ly's "Synonymie Catalogue" (1889).
RlTCHIE, 1911, has given a recent synonymy of this common and widely distributed species; which makes it unnecessary for me to do
more than refer to the original description and to those notices of the species which are important, in the present connexion, from the
point of view of Geographical Distribution, or in other ways.
27
28
Barentsia gracilis Annandale, 1912, "Occ. Ent. hul. Waters'*, Ree. Ind. Mus.. VII, p. 205
(Madras).
Barentsia gracilis Waters, 19 14, "Mar. Fauna Brit. E. Air.", Proc. Zool. Soc, p. S55.
opodaria gracilis Kirkpatrick, 1888, "Pol. Port Phillip", Arm. Mag. Nat. Hist. (6) II, p. 21
(Victoria ').
Ascopodaria gracilis Ehlers, 1890, "Kenntn. Pedicellineen", Abh. Göttingen, XXXVI, p. 142.
Ascopodaria gracilis Robertson, 1900, "Stud. Pac. Coast Ent.", Proc. Cal. Acad. Sci. (3) Zool.
II, p. 345 (California).
Pedicellina gracilis, var. nodosa Lomas, 1886, "Rep. Pol. L. M. B. C. Distr.", Proc. Lit. Phil.
Soc. Liverpool, XL, p. 190, PI. III, fig. 2 (Isle of Man; the variety was distinguished by
the nodes of the stalk).
Ascopodaria nodosa Lomas, 1887, "On Asc. nodosa'', Ibid., XLI, p. XLVI.
Ascopodaria nodosa Lomas, 1889, "Second Rep. Pol. L. M. B. C. Distr.", Proc. Biol. Soc.
Liverpool, III, p. 217.
Barentsia nodosa Duerden, 1S93, "New and rare Irish Pol."', Proc. R. Irish Acad. (3) III,
p. 135, PI. V, fig. 4.
Gonypodaria nodosa Ehlers, 1890, t. cit, p. 144.
433. B. Stat. 7. Reef of Batjulmati, Java, reef; coral and stones (on 2. A., Scrupocellaria ferox).
20. J. Stat. 50. Bay of Badjo, W. coast of Flores, o — 40 Metres; mud, sand and shells (on
20. I., Scrupocellaria).
24. F.) Stat. 53. Bay of Nangamessi, Sumba, o — 36 Metres ; coral-sand; near the shore, mud
31.J. J (on 24. C.:', Mimosella tennis; and on 31. I.2, Scrupocellaria).
427. D. Stat. 60 and 303. Haingsisi, Samau Island, Timor, o — 36 Metres; Lithothamnion (on
427. B.\ Catenaria).
337. H.) Stat. 71. Makassar, o — 32 Metres; mud, sand with mud, coral (on 337. A., Scrupo-
338. D.) cellaria; and on a Hydroid on slide 338. C, Valkeria atlanticd).
37.ZA.J1 Stat. jj. Borneo Bank, 59 Metres; fine grey coral-sand (on 3/. O., Canda; and on
41. C. | 41. A., Bugula).
108. AH. Stat. 144. Anchorage North of Salomakiëe (Damar) Island, 45 Metres; coral-bottom
and Lithothamnion (on 108. AE., Beania).
376. P. Stat. 213. Saleyer, o — 26 Metres; coral-reefs, mud and mud with sand (on 376. C,
Schizoporella).
425. C. Banda Sea (on sea-weed, 011 slide 425. B., Aeted).
296. E. Stat. 273. Anchorage off Pulu Jedan, Aru Islands, 13 Metres; sand and shells (on
296. B.1, Catenicella).
259. G.) Stat. 318. 6° 36'. 5 S., ii4°55.5E., 88 Metres; fine yellowish grey mud (on 259. C,
415. B.) Carbasea cribriformis ; and on 415. A., Flus/ra re/iculiuii).
262. F. Stat. 319. 6°i6'.5S., U4°37'E., 82 Metres; fine yellowish grey mud (on 262. L.,
Flus tra reticu lit m ) .
555. K. Stat. 320. 6°5'S., II4°7 E., 82 Metres; fine grey mud (on 555. C, Bugula, mounted
on slide 555. J.~, Lagenipora).
Singapore, Mus. Zool. Cambr., Reg. Nov. 11, 1899, F. P. BEDFORD Coll., 5 fathoms.
Japan, Mus. Zool. Cambr., Reg. June 23, 1902. A. OWSTON Coll., near Tokyo, 30 — 40 fathoms.
Size smal], usually less than 1 mm. in total length (stalk + calyx). Thin part of the
stalk variable in length, not infrequently showing an incomplete, or more rarely a complete
muscular node towards its distal end, or near the middle. Tentacles apparently about 14.
The present species has been found, in the majority of cases, in small quantities, on
I) In Mr KlRKPATRICK's specimens fiom Port Phillip, in the Collection of the British Museum (SS. 5. 17.20 and 22) I do not
find that the distal end of the chitinous part of the stalk is pointed like the corresponding part of PediccUinopsis fntticosa, as described
by that author in the paper cited.
2S
29
slides which have been mounted to illustrate other species of Polyzoa. It is obvious that it is
widely distributed in the Malay Archipelago; and I have no doubt that it might be found on
Hydroids, Polyzoa, etc, from many Stations from which I have not recorded it.
The 'Siboga' specimens agree closely witli those from Europe. The individual shown by
Sars in fig. 2Ó of the original description resembles them in having the indication of a muscular
joint on the stem. In one specimen (337. H) I have found one complete stalk-joint, and a
second, more distally placed joint which is not completely formed. Other examples of jointed
specimens of this species are noted in the synonymy. The lengths, in u, of the several parts of
the individual shown in fig. 12 are as follows: — calyx, 190; thin part of stalk, 420 ; muscular
base, 150; total length, 760 ; breadth of calyx, 180. The thin part of the stalk varies greatly
in length in the specimens examined. I have not been able to ascertain the number of the
tentacles with certainty, but it seems to be about 14. In the specimens 338. D., but not in
other cases, I have noticed a few short, curved spines on the calyx, on the soft piece of the
stalk next the calyx, and on the stem-joint, if this is present. Waters (19 10, p. 282) records
similar spines in specimens of this species from the Sudanese Red Sea, Trieste and the South
coast of England. Most of the specimens seem to be sexually immature.
The present species differs conspicuously from others obtained by the 'Siboga' in its
delicate character and much smaller size.
2. Barentsia discreta Busk. (PI. II, figs 8, 9).
Ascopodaria discreta Busk, 1886, "Challenger Reports", L, "Polyzoa", Pt II, p. 44, PI. X,
figs 6 — 12 (Tristan da Cunha, 100 — 150 fathoms).
Ascopodaria discreta Ehlers, 1890, "Kenntn. Pedicellineen", Abh. Göttingen, XXXVI, p. 143.
Ascopodaria discreta Thornely, 1905, HERDMAN's "Rep. Pearl Oyster Fisheries", Publ. by the
Roy. Soc, Suppl. Rep. XXVI, p. 128 (Ceylon).
Barentsia discreta Kirkpatrick, 1890, "Hydr. Pol. China Sea", Ann. Mag. Xat. Hist. (6) V,
p. 17 (Tizard Bank, 27 fathoms).
Barentsia discreta Waters, 1904, "Rés. Voy. Belgica", Zool., "Bryozoa**, p. 99, PI. VII,
figs 6a — ■ƒ; PI. VIII, figs \-ja — c (Ile Londonderry, Magellanes Chili).
Barentsia discreta Waters, 1905, "Bry. Cape Horn", J. Linn. Soc, Zool., XXIX, p. 230.
Barentsia discreta Osburn, 1910, "Bry. Woods Hole". Buil. Bur. Fisheries (Washington), XXX,
p. 214, PI. XVIII, figs 5, $a (Woods Hole Region, U. S. A.; Florida).
Barentsia discreta Annandale, 19 12, "Occ. Ent. Ind. Waters", Ree. Ind. Mus. VII, p. 205
(Port Canning, India). The reference to this species does not seem certain.
Barentsia discreta Sumner, Osburn and Cole, 1913, "Biol. Surv. Woods Hole", Buil. Bur.
Fisheries, XXX, Pt I, p. 108; Pt II, p. 596.
Barentsia discreta Osburn, 1914, "Bry. Tortugas Islands", Publ. 182, Carnegie Inst. Washington,
p. 1S5 (Florida).
Ascopodaria misakiensis Oka, 1890, Zool. Mag. N" 20 (memoir not seen).
Barentsia misakiensis Oka, 1895, "Sur B. misakiensis", Zool. Mag., VII, N" 78, p. 76, PI. XII,
figs 1 — 8 (S. of Tokyo).
Waters (1905) gives Pedieellina australis Jullien, 1888 ("Miss. Sci. Cap Horn", VI,
"Bryozoaires", p. 13, PI. VIII, fig. 4) {nee Ridley, 1881) as a synonym of the present species;
and Osburn, 19 14, regards B. timida Verrill 1899 — 1900 ("Tunicata and Molluscoidea of the
Bermudas", Trans. Conn. Acad., X, p. 594) as being probably another svnonym.
3Q
195- B. Stat. 274. 5°28'.2S., i34°53'.9E., 57 Metres; sand and shells, stones (on Retepora,
195. A.).
131. S. Stat. 164. i°42'.5 S., I30°47'.5 E., 32 Metres; sand, small stones and shells (on Lepralia
celleporoideSi 131. A.).
337. I. Stat. 71. Makassar, O — 32 Metres; mud, sand with mud, coral.
348. A. Stat. 213. Saleyer, o — 36 Metres; coral-reefs, mud, and mud with sand (on Adeonella,
347. A.).
Size large, reaching a total length (calyx + stalk) of at least 3 mm. Tentacles about
20 — 24. Stalk with a large proximal muscular portion and a very long, cylindrical, distal
portion, without spines but covered with a thick cuticle, the inner layer of which is perforated
by numerous pores.
If the synonymy given above is correct, B. discreta has a wide distribution. The specimens
originally described by Busk were dredged off Tristan da Cunha, 100 — 150 fathoms (Brit. Mus.
99. 7. 1. 4447). It has been recorded more recently by Kirkpatrick from the Tizard Bank, China
Sea, 27 fathoms (Brit. Mus. 89. S. 21.66); by Miss Thornely from Ceylon; by Annandale
from India; by Oka from near Tokyo, 5 — 10 Metres; by Waters from 11e Londonderry, Canal
Francais, Magellanes, Chili; by Osburn from Woods Hole, Mass. and from Florida; and, assuming
the correctness of Waters' synonymy, by Jüllien from Ile Hoste (near Cape Horn), 26 Metres;
and perhaps by Verrill from the Bermudas. The 'Siboga' localities fall into two groups; -
Stat. 274 and Stat. 164 from near the N. W. extremity of New Guinea; and Stat. 71 and
Stat. 2 1 3 from the S. end of the Straits of Makassar and the neighbourhood.
I have no doubt that the 'Siboga' specimens belong to the same species as the forms
described by Oka under the name of Barcntsia misakiensis. The original description given by
that author appeared in Japanese; and I derive my knowledge of his results from his later
paper (1895), which was published in French. I have been able to verify his account of the
Japanese form by the examination of specimens obtained from near Tokyo, 40 — 53 fathoms, in
the University Museum of Zoology, Cambridge (Reg. June 23, 1902). Although I have counted
23 tentacles in one of the Japanese individuals, coming within the limits (20 — 24) given by
Oka, another was found to have as many as 29. Busk gives 16 — 20 for the specimens from
Tristan da Cunha. But independently of the fact that the range of variation in the number of
tentacles in Pedicellinidae has not been satisfactorily decided, too much stress must not be laid
on statements with regard to the number, unless the author in question has had specially good
opportunities of counting them with certainty. I am thus not disposed to attach too much
importance to Oka's statement (1895, p. 5) that B. misakiensis can easily be distinguished from
B. discreta by the larger number of its tentacles.
The present species appears to be well characterised by its large size; by the great
length of the narrow portion of the stalk; and, above all, by the curious pores with which this
part of the stalk is provided. These pores may perhaps be regarded as vestigial spines. Similar
pores are known to occur in the stalks of certain other Pedicellinidae, as in Pedicellinopsis
tru/icosa Hincks, in Barentsia macropiis Ehlers and in B. (Gonypodaria) ramosa Robertson.
In all these respects the 'Siboga' specimens agree closely with those from Japan. This
may be illustrated, with regard to size, by the following measurements, in a :
3°
Length of calyx .
Length of narrovv
part of stalk . .
Length of basal
part of stalk . .
Total length . . .
'Siboga', 195. 11.
Japan.
'Siboga', 195. B.
Japan.
432—688
448—752
Breadth of calyx .
Breadth of narrow
384—460
44O — 560
880 — 2224
lÓOO — 2176
part of stalk . .
Breadth of basal
45
45
2/2—430
432— S7r>
part of stalk . .
140
192
I584 — 3OOO
2480 — 3104
It will be seen that some of these measurements are variable, the size of the calyx,
however, depending on its age ; and the dimensions of the basal muscular part of the stalk
probably depending on its state of contraction. The Japanese specimens have, moreover, the
appearance of being less contracted than those obtained by the 'Siboga'. But there can be no
question that the narrow part of the stalk is very variable in length ; the cuticle being so thick
that the question of contraction need not be considered. The variability in length appears,
however, to be mainly the result of differences in growth. The individuals near the growing
points have short stalks ; and it is clear, from a comparison of these with older specimens,
that growth continues for a long time in the neighbourhood of the calyx, where the cuticle
disappears. Oka has called attention to the fact that the calyces are deciduous and are readily
regenerated ; a property which this species shares with other Pedicellinidae : — for instance
Pedicellina cernua l). In the Japanese specimens I find that this process may be associated with
an increase in the length of the stalk, a distinct annular mark being visible at the junction of
the old and of the regenerated portions. The part thus added is very variable in length, but
it may be as long as the original stalk. Sometimes tvvo or even three annular marks are
visible, indicating as many successive regenerations. In the specimen figured (fig. 9) there is
some alteration of the cuticle near the middle of the stalk, suggesting that a joint might have
been formed later here ; and in another specimen a joint, with indications of the development
of longitudinal muscles, has actually been formed in a corresponding region. Specimens of B.
discreta with a jointed stalk have been figured by Waters (1904, PI. VII, figs 6a, 6d),
The peculiar pores of the thinner part of the stalk traverse the inner layer of the cuticle,
but are closed by an external layer which, according to Oka, differs from the inner stratum
by staining with eosin. The position of the pore is sometimes, but not always, indicated by a
very slight convexity of the outer surface of the stalk. The basal, muscular portion of the stalk
is usually not separated from the thinner part by a diaphragm, although some of the Japanese
specimens show an annular thickening of the chitin, projecting slightly into the cavity of the
stalk, in that position. In one of the 'Siboga' specimens (337.1) the diaphragm is well developed,
and the longitudinal muscles of the basal thickening of the stalk are inserted into it.
The stolon is composed of alternating segments, divided from one another by discoidal
diaphragms perforated at the middle. Each alternate segment gives off a stalk, the intervening
segments being barren. This arrangement may be considered typical in the Pedicellinidae and
has been correctly described by Oka for the present species. A lateral branch of the stolon is
1) Cf. my paper on that species in Quart. J. Mier. Sci., XXVII, 1887, p. 256.
31
32
commonly given off, at right angles, on each side of the polypiferous segments (fig. 8). In one
case (195. B.s) two branches are given off on one side and one on the other.
I have noticed two slight differences between the 'Siboga' specimens and those from
Japan. In the latter there is a long cylindrical oesophagus, and the stomach has a conspicuous
lobe projecting from its ventral (lophophoral) side towards the oesophagus. In the 'Siboga'
specimens the cylindrical part of the oesophagus is short, while the lobe of the stomach is not
present, although it is perhaps represented by a slight indentation in the inner surface of the
epithelium, in the position of the lobe of the Japanese specimens. These differences do not
seem to warrant a separation of the two forms, in view of the close resemblance which exists
in other respects.
Oka states that the sexes are separate and that the colonies are unisexual.
The present species seems to be nearly allied to B. macropzis Ehlers, 1890 and to
B. timida Verrill. The latter name is regarded as probably a synonym of B. discreta by
Osburn, 19 14.
o
. Barentsia laxa Kirkp. (PI. II, figs 10, 11).
Barentsia laxa Kirkpatrick, 1890, "Hydr. Pol. Torres Straits", Proc. R. Dublin Soc. (N. S.)
VI, p. 624, PI. XVII, fig. 6.
88. A. Stat. 115. East side of Pajunga Island, Kwandang Bay, reef. On a Coral {Porties).
380. D. Stat. 47. Bay of Bima, 55 Metres; mud with patches of fine coral-sand.
Size large, reaching a total length of nearly 3.5 mm. Basal muscular part of stalk either
without appendages or bearing a pair of distally situated, pointed, membranous appendages.
Thinner part of stalk at first cylindrical, then increasing in width and annulated with numerous
rings; this part of the stalk very flexible.
The characters of the stalk are markedly different from those in the previous species,
in which the greater part of the length is rendered rigid by a strong development of the cuticle.
In B. laxa it is obvious that the contractility is not confined to the basal portion, but is also
present in the greater part of the length of the stalk. This is shown by the extremely variable
shape of the preserved specimens, in which the annulated part of the stalk is curved into all sorts
of positions1); and is also indicated by the fact that longitudinal muscles can be distinguished
in this region.
The measurements of the specimen represented in fig. 1 1 are as follows (in u.) :
Length of calyx 480 Breadth of calyx 365
Length of distal part of stalk .... 2500 Breadth of stalk, near calyx 80
Length of muscular base of stalk . . 480 Breadth of stalk, near muscular base. 32
Total length 3460 Breadth of muscular base 190
1) In the contractile character of the distal part of the stalk this species resembles a form of Barentsia from Newport, R. I.,
U.S. America, described by I.eidy (" Urnatella gracilis", J. Acad. Nat. Sci. Philadelphia, (2) IX, 1S84, p. 14, text-fig. on p. 11), and
on idered by him to be nearly allied to H. gracilis. The stalk of B. major Hincks ("Pol. St Lawrence", I, Ann. Mag. Nat. Hist. (6) I,
iSSS, p. 226, PI. XV, fig. 2) and of its var. elongata Jullien and Calvet (1903, p. 27, PI. II, fig. 3) is similarly annulated. In B. variabilis
Calvet and in B. capitata Calvet (1904, „Kry. Hamburg. Magalhaensische Sammelreise", pp, 40, 41, PI. III, figs I, 2), a stiff, chitinous
part of the stalk is suceceded distally by an annulated, fiexible part.
32
oó
The calyx is ovoid in form, and the alimentary canal lies in close contact with the
body-wall. The number of the tentacles has not been counted with certainty, bnt it appears to
be about 20. The two rounded boclies seen in fig. 1 1 close to the intestine are gonads, the
one nearer to the intestine being almost certainly an ovary, and the more orally situated body
having the appearance of a testis. If this observation be correct the species is hermaphrodite.
There has been considerable difference of opinion among authors with regard to the
question whether the Pedicellinidae are hermaphrodite or dioecions ; and it seems probablc that
both conditions really occnr in the Family. In the well known account given by Nitsche, the
hermaphrodite condition is clearly figured1); but in this case the ovaries are on the oral sidc
of the testes. If my account is correct the opposite condition appears to be characteristic of
the present species, in which, moreover, the gonads lie close to the intestine, instead of being
separated from it by h large brood-pouch. But as the gonads are clearly young in fig. 11 it is
not impossible that their position might have been shifted later by the development of a brood-
pouch from the floor of the vestibule.
The stolon appears to be quite normal in its characters; and a branch may be given
off from the base of a stalk (fig. 10).
Regeneration of the calyces is indicated by their very varying size in the specimens 88. A. ;
a large stalk often bearing a very small calyx.
The measurements given above refer to N° 88. A., in which the muscular base of the
stalk bears no distal appendages. In 380. D. the stalk is not more than one third the length of
that shown in fig. 1 1 (88. A.), but is not easy to measure in consecjuence of the curved attitude
it has assumed. From the observations above recorded on D. discreta it is not improbable
that the very long stalks of 88. A. may have been the result of regeneration of the calyces.
The specimens 380. D. are, however, characterised by the constant possession of a pair of
membranous appendages, attached to the distal end of the muscular base of the stalk ; the
characters of which are so similar, in other respects, to those of 88. A. that it does not seem
to be necessary to refer the two sets of specimens to different species.
The determination of the 'Siboga' specimens has been verified by comparison with the
typ'e-specimens (Torres Straits, 6 — 7 fathoms; 90. 11.22. 1.) in the British Museum. The types
agree closely with 88. A. in size, in the absence of membranous appendages on the muscular
base, and in the flexible character of the distal part of the stalk, which, moreover, shows traces
of the 'annulated' character noticed in the 'Siboga' specimens. The total length, as recorded
by Kirkpatrick, is 3000 fji; the calyx being 400 — 500 u. in length and 350 — 400 u. in breadth.
It may be remarked that the distal part of the stalk in the original figure of this species is
represented in a condition shrivelled by the action of reagents.
4. Bare?itsia geniculata n. sp. (PI. II, figs 6, 7).
Type. 260. E. Stat. 318. 6° 36.5 S., II4°55'.5E., 88 Metres; fine yellowish grey mud (on 260. A.,
Stichoporina simplex).
1) Nitsche, H., 1870, ("Üb. d. Anat. v. Pedicellina tchinata", Zeitsclir. wiss. Zool., XX, p. 26, PI. II, fig. 2. D., PI. III, fig. 5.
33
S1BOGA-EXPEDITIE XXVIII (7. 5
34
2Ó2. H. Stat. 319. 6° 16.5 S., 1140 37' E., 82 Metres; fine yellowish grey mud (on 282. B.,
Stichoporina simplex).
Size large, reaching a total length of about 2.85 mm. Stalk interrupted by two or three
muscular joints, the intervening parts covered by a well developed cuticle, which is smooth or
has only the barest traces of tubercles. These internodes are cylindrical in form, expanding as
they reach the joints. Tentacles probably not less than 20 — 24.
I am unable to refer this species to any of the forms of Barentsia previously known.
In general appearance it has considerable resemblance to the jointed forms of B. discrcta
described by Waters l) from Ile Londonderry, Chili. Waters gives Pedicellina australis Jullien
{11011 Ridlev) as a synonym of B. discrcta ; and a reference to Jullien's account') shows that
in the form described by that author the stalk may be jointed. It is not impossible that the
specimens here described may be identical with Jullien's species, although> it appears improbable
that they are jointed forms of B. discrcta, since the pores so characteristic of the stalk of that
species are not present.
The muscular base of the stalk is divided from the next part by a chitinous diaphragm.
Waters has described a similar arrangement in his Chilian specimens; but in those which I
have referred to B. discrcta the diaphragm is usually not present, or is represented (in Japanese
specimens) by a slight annular thickening of the cuticle, which hardly projects into the cavity
of the stalk.
A similar diaphragm occurs on the distal side of each of the stem-joints; and, as Waters
has suggested, it perhaps serves for the insertion of the longitudinal muscles of the basal
dilatation and of the joints. Evidence of the insertion of muscles into a diaphragm has been
alluded to in the description of B. discrcta (see p. 31).
The following are the lengths, in u, of the several parts of the individual shown in
fig. 6: — basal dilatation, 250 ; first internode of stalk, from the proximal diaphragm to the
commencement of the joint, 860 ; first joint, to its distal diaphragm, 150; second internode oi
stalk, 5 80; second joint, 130; third internode, 270; calyx, 400 ; total, 2640. The breadth of
the calyx is 310 p..
I am not able to state the number of the tentacles with certainty, but it appears to be
about 24. The specimen figured seems to be a female, with an embryo in its brood-pouch.
The stem-internodes of this species are much longer than those of B. (Arthropodaria)
benedeni Foettinger. They more nearly resemble those of a specimen referred by Ritchie s) to
B. gracilis, which is, however, a much smaller species than the one here described. The length
(in the figure) of the longest individual shown by Ritchie in fig. 5 is about 115 mm. As the
figure is said to be X l4°, this gives a total absolute length of less than 1 mm.
1) Waters, A. W., 1904, p. 99, PI. VII, figs 60, 6</.
2) Jullien, J., 188S, p. 13, PI. VIII, fig. 4.
3) RITCHIE, J., 191 1, Trans. R. Soc. Edinburgh, XLVII, Pt 4, p. S41.
34
35
Sub-Class II. ECTOPROCTA Nitsche.
1869, Zeitschr. wiss. Zool., XX, p. 596.
Order I. GYMNOLAEMATA Allman.
1S56, "Mort. Fresh-water Pol.", p. 10.
Sub-Order I. CTEXOSTOMATA Busk.
1S52, J. MacGillivray's "Voy. Rattlesnake", I, p. 346.
Halcyonellea Ehrenberg (pars), 1839, Üb. d. Bild. cl. Kreidefelsen", Phys. Abh. k. Ak. Wiss.
Berlin, A. d. Jahre 1838,- Tab. II.
Group A. Carnosa Gray.
1841, "Syn. Brit. Mus."', 43rd Ed., p. 149.
1848, ''List Brit. Animals Brit Mus.", I, pp. 91, 144.
Alcyonellea Hincks, 1SS0, "Hist. Brit. Mar. Pol.", p. 490.
Alcyonellea Waters, 1910, "Rep. Mar. Biol. Sudanese Red Sea", "Bryozoa II", J. Linn. Soc.
Zool., XXXI, p. 240.
The present group includes what may be termed the encrusting forms of Ctenostomata.
For this series Hincks has used the name "Halcyonellea Ehrenberg", placing in it the Families
Alcyonidiidae, Flustrellidae and Arachnidiidae. The name Halcyonellea had been previously
employed, in a more or less similar sense, by other writers, as by Johnston :). But a reference
to the original introduction of the term will show that Ehrenberg3) gives "Halcyonellea.
Federbusch-Polypen. Genera: Halcyonella et Sertularinorum reliquiae". Taking into
consideration Ehrenberg's earlier account 3), where Alcyonclla includes A. stagnorum (= A.
tungosa auctt.) and A. articulata (= Paludicella)y it follows that the name "Halcyonellea",
as originally introduced, referred to the Phylactolaemata as much as to the Ctenostomata.
In Ehrenberg's later and more complete classification (1839, 1. c), Halcyonellea included
Halcyonella and various Ctenostomata ( Vesicularia, Boweröaiikia, Farrella, Valkeria and
Alcyonidiitm), besides Stephanidiitm [a Hydroid ?] and Dynaniene [a Hydroid]. In view of
these facts I employ Gray's name, which comprised in the text of the work (1848) two Families
only : — the Alcyonidiadae, consisting, vvith one exception, of species of Alcyonidituu^ and the
Pedicellinidae. The latter are, however, transferred to the order Cornea in the Appendix
(p. 145) of the same Catalogue, so that the Carnosa were there left by Gray with the single
Family Alcyonidiadae.
ij Johnston, G., 1847, p. 358.
2) Ehrenberg, C. G., 1834, "Die Corallenthiere d. rothen Meeres", Abh. k. Akad. Wiss. Berlin, p. 153, sep.
3) Ibid., 1831, "Symbolae Physicae", "An. Evert.", Dec. I, "Phytozoa Polypi", sheet a.
35
36
Fam. Alcvonidiidak Johnston.
Alcyonidiêes (part) Lamouroux, 1813, Ann. Mus. Hist. Nat. XX, p. 2S4.
Alcyonidulae Johnston, 1838, "Hist. Brit. Zooph.", Ed. i, p. 300.
Alcyonidulae Couch, 1844, "Cornish Fauna", III, p. 132.
Alcyonidiadae Johnston, 1837, Trans. Benvicksh. Nat. Club, I, p. 108.
Alcyonidiadae Gray, 1848, "List Brit. An. Brit. Mus.", I, pp, 91, 144.
Alcyonidiidae Hincks, 1S80, "Hist. Brit. Mar. Pol.", p. 490.
Alcyonidium Lamx.
Alcyonidium Lamouroux, 1813, "Essai Gen. Thalassiophytes" (Suite). Ann. Mus. Hist. Nat.
(Paris) XX, p. 285.
Alcyonidium Lamouroux, 1821, "Exp. Méthod.", p. 71.
Alcyonidium was introduced by Lamouroux (18 13) as a genus of Sea-weeds. The genotype
is A. diaphanum = A. gelatinosum L.
No specimens of Alcyonidium have been found among the 'Siboga' dredgings; but I
include the genus in the present Report on the evidence of a single specimen from Torres Straits.
As Waters l) has pointed out, Alcyonidium has rarely been recorded from the Southern
Hemisphere. Besides A. antarcticum, described in the Memoir here referred to, Waters mentions
A. Jïaöclliforme Kirkpatrick, 1902 (Antarctic), A. flustroides Busk, 1886 (Cape of Good Hope)
and A. gelatinosuni, mentioned by Hincks2) as having been found off the coast of Natal. In
the absence of any evidence on the subject, some doubt may be expressed with regard to the
correctness of the last determination. To the list given by Waters may, however, be added
A. mytili, recorded by Kirkpatrick (1888) from Victoria and A. mytili Calvet from Tierra
del Fuego. From Oriental waters Miss Thornely has given A. mytili as occurring in the Gulf
of Manaar. These records are quoted in the subjoined synonymy. From the Northern Pacific
Miss Robertson 3) has described four species of the genus, three of which are determined as
belonging to British species. In a later paper +) the same author has described another species,
A. pedunculatum, besides giving a new description of the remarkable Ascorhiza occidentalis
Fewkes, 1889, also referred to the Alcyonidiidae. It has, however, been pointed out by Waters ')
that Ascoi'hiza is synonymous with Clavopora^ described by Busk ü) in 1874 from the dredgings
of the 'Porcupine' in the Mediterranean, in deep water off the African coast.
It may be concluded from these facts that the Alcyonidiidae are mainly Northern forms,
and specially Atlantic. In other parts of the world they would appear to be most common in
the North Pacific. As the genus Alcyonidium extends into high latitudes in the Atlantic area,
its occurrence elsewhere may have been due to its invasion of the Pacific from the North.
il WATERS, A. W., 1904, "Kés. Belgica", "Bryozoa", p. 86; see also 1904, Min. Fr.-Josef Land", II, J. Linn. Soc. Zool.,
XX1N, p. 178.
2) Hincks, T., 1880, p. 492.
3) Robertsox, A., 1900, "Harriman Alaska Exp.", "Bryozoa", Proc. Washington Acad. Sci.. II, pp. 329, 330.
4) Ibid., 1902, "Obs. on Ascorhiza.. and Related Alcyonidia", Proc. Calif. Acad. Sci., (3) Zool., III, pp. 104 —
5) Waters, A. W., 1910, p. 240.
6) BüSK, <... 1874, "C/avopora hystricis", Quart. J. Mier. Sci., (N. S.) XIV, p. 261.
36
37
i. Alcyonidium polyoum Hassall. (PI. III, fig. i).
Sarcochitum polyoum Hassall, 1841, "Suppl. Cat. Irish Zooph.", Ann. Mag. Nat. Hist. VII, p. 484.
Alcyonidium mytili Dalyell, 1848, "Rare and Remark. An. Scotland", II, p. 36, PI. XI.
figs 1 — 14.
Alcyonidium mytili Hincks, 1880, "Hist. Brit. Mar. Pol.", p. 49S, PI. LXX, figs 2, 3.
Alcyonidium mytili Levinsen, 1894, "Zool. Danica", IX, "Mosdyr", p. 81, PI. VII, figs 33 — 37.
Alcyonidium mytili Silbermann, 1906, uUnt. Alcyonidium mytili", Arch. f. Naturg., Jahrg. 72,
Bd I, p. 265, Pis XIX, XX.
Alcyonidium mytili Osburn, 1912, "Bry. Woods Hole", Buil. Bur. Fisheries, XXX, p. 251,
PI. XXVIII, figs 74, 740.
Alcyonidium mytili Sumner, Osburn and Colc, 191 3, "Biol. Surv. Woods Hole", Buil. Bur.
Fisheries, XXXI, Pt I, pp. 108, 109; Pt II, p. 606.
The following records, without description or figure, may also be noticed :
Alcyonidium mytili Robertson, 1900, "Bry. Harriman Alaska Exp.", Proc. Washington Acad.
Sci., II, p. 329.
Alcyonidium mytili Calvet, 1904, "Bryozoen", Hamburg. Magalhaensische Sammelreise, p. 38.
Alcyonidium mytili Thornely, 1905, Herdman's "Rep. Pearl Oyster Fisheries Gulf of Manaar",
Publ. by the Roy. Soc, Suppl. Rep. XXVI, p. 127.
Kirkpatrick's A. mytili, from Victoria ("Pol. Port Phillip", Ann. Mag. Nat. Hist., (6)
II, 1888, p. 17, PI. II, figs 6, 6a) does not appear to me to belong to this species.
It is probable that Sarcochitum polyoum Hassall, 1841, is the form assumed by old
colonies of A. mytili. If this be the case, Hassall's name has the priority, as has been pointed
out by Osburn (191 2). Duerden *) has recorded A. mytili from Dublin Bay, and considers that
it is identical with Hassall's species, from the same locality. Silbermann (1906, p. 266), in a
recent study of A mytili, comes to the conclusion, however, that this species is distinct from
A. polyoum.
Murray Islands, Torres Straits, 15 — 20 fathoms, on Euthyris obtecta Hincks. Mus. Zool.
Cambridge, Reg. Sept. 1, 1900; now transferred to the British Museum.
Zoarium forming a thin film, the interzooecial septa very distinct. Zoocia more or less hexa-
gonal, averaging about 450^ in length ; the orifice on a low papilla which is not quite terminal.
I do not teel certain that the present form is correctly referred to the European species ;
but there is nothing in the characters given above to forbid this association.
The single specimen under consideration forms an extremely thin, transparent film
encrusting a part of the frontal surface of a colony of Euthyris obtecta. lts interzooecial septa,
which are very distinct, have been developed without any relation to those of the Cheilostome
on which it is growing. Some of them cross the opercula of the Euthyris, the corresponding
polypides of which have degenerated, although they are present beyond the growing margin
of the Alcyonidium : - - indicating that the degeneration has been the result of the occlusion
of the orifices.
The orifice of the Alcyonidium is of the usual type, being ver)- small and circular, or
transversely oval, in the retracted condition, and being situated at the summit of a low papilla,
1) Duerdex, J. E., 1894, "Notes Mar. Idv. Rush, Co. Dublin'', Irish Nat., III, p. 232.
37
33
vvhich is not quite terminal. The tentacles are about 12 in number. The tentacle-sheath may
lie either to the right or to the left of the intestinal limb of the alimentary canal; no uniformity
in this respect being discoverable, even in the zooecia of a single longitudinal row. The pharynx
is short; and in the retracted condition its proximal end forms the extremity of the first limb
of the alimentary canal. The oesophagus, formed of transparent cells, is long and passes distally.
It is remarkable for its width l), which exceeds that of the caecum of the stomach. The stomach
and its caecum are noticeably small, and together do not greatly exceed the length of the
rectum, which is well developed. The parietal muscles are small and delicate, the fibres occurring
singly. The retractor-muscles form a conical group, embracing the proximal end of the lophophore.
Reproductive organs are absent.
Waters 2) has given a useful list of the number of tentacles believed to be characteristic
of various species of Alcyonidium. None of these records are so low as 1 2 except those of
Lamouroux and Fleming (both referring to A. gelatinosum; which, according to Hincks, has
15 — 17 tentacles); and Waters considers that the number 12 has been incorrectly recorded
by those authors. The number given for A. viytili is 15 — iS, on Hincks' authority, while
Silbermann 3) gives 16 — 20. It is difficult to count the retracted tentacles of a Polyzoon seen
in frontal view, in an entire preparation ; but in the case of the Torres Straits specimen the
zooecia are so transparent that the probable error of my enumeration is not very great.
As in other cases, there is some irregularity in the size and shape of the zooecia, which
are, however, fairly regularly hexagonal. They average about 400 u. in length and 290 <x in
breadth. The longest and narrowest measured is 560 by 225 u; and the broadest zooecium is
400 by 385 u..
It should not be concluded, without further evidence, that the thinness of the zoarium is
really characteristic of the Torres Straits form. A. polyoum from European waters is well known
to increase in thickness during growth, the central parts of the colony thus acquiring a very
different appearance from that of the delicate marginal regions (cf. Silbermann, 1906, p. 268).
Fam. Flustrellidae Hincks.
Flustrellidae Hincks, 1880, ''Hist. Brit. Mar. Pol.", p. 504.
Elzerina Lamx.
Eiser ina Lamouroux, 18 16, "Hist. Pol. Cor. Flex.", p. 122.
1. Elzerina blainvillii Lamx. (PI. III, fig. 2).
Elzerina blainvillii Lamx, 18 16, t. c, p. 123, PI. II, figs 3 a, 3 B.
Elzerina blainvillii Lamx, 1821, "Exp. Méthod.", p. 3, PI. 64, figs 15, 16.
i) A similar character seems to be indicated in the form from Woods Hole, referred to A.mytili by OSBURN (1912, PI. XXVIII,
fig. 74»); and it is also shovvn by Silbermann (1906, PI. XIX, figs 1 — 4) in specimens from the Baltic.
2) Waters, A. W., 1904, "Bry. Franz-Josef Land" II, J. Linn. Soc, Zool., XXIX, p. 180.
3) Silbermann, 1906, t. cit., p. 272, PI. XIX, figs 6, 7.
38
Elzerina blainvillii Lamx, 1824, "Encycl. Méthod.", "Zoophytes", p. 317.
Elzèrina blainvillii Blainville, 1S34, "Man. Actinol.", p. 453, PI. LXXX, figs 2, 2 a.
Elzerina blainvillii Milne Edvvards, 1836, LAMARCK's "Hist. An. s. Vert.", 2 Ed., II, p. 240.
Elzerina blainvillii Gray, 1843, DlEFFENBACH's uNe\v Zealand", II, p. 293.
Verrucularia dichotoma v. Suhr, 1834, Flora, XVII. Jahrg., I. Bd, p. 725, PI. I, figs 9, a, b.
Verrucularia dichotoma Busk, 1SS4, Challenger Report, Pt XXX, p. 48.
Verrucularia dichotoma MacGillivray, 1887, "Cat. Mar. Pol. Victoria", Trans. R. Soc. Vict.i
XXIII, pp. 189, 205 (Sep., 3, 19).
Verrucularia dichotoma MacGillivray, 1890, McCov's 'Trodr. Zool. Vict.", II, Dec. XX,
p. 348, PI. 195, figs 6— ja.
Verrucularia dichotoma Whitelegge, 1890, "List Mar. F.-W. Fauna Port Jackson", J. Proc.
R. Soc. N. S. Wales, XXIII, p. 123.
Farciminaria dichotoma Busk, 1S61, Quart. J. Mier. Sci., (N. S.), I, p. 155, PI. XXXV,
figs 1 — 1 b.
Farciminaria blainvillii Hutton, 1891, "Rev. List Mar. Bry. X. Zealand", Trans. Proc. X. /..
Inst., XXIII, p. 104.
Flustrella dichotoma Hincks, 1884, Ann. Mag. Nat. Hist. (5) XIII, p. 366, PI. XIV, figs 2— 2b.
Flustrella dichotoma Hincks, 1893, Ibid-, (6) XII, p. 141.
Torres Straits, A. C. Haddon Coll., Mus. Zool. Cambridge, Reg. Feb. 24, 1898; now trans-
ferred to the British Museum.
Timor , Australasia (Lamouroux) ; Algoa Bay (v. Suhr) ; \Tictoria (Busk: , Hincks ,
MacGillivray); Port Jackson, Sydney (Whitelkgge and Mus. Zool. Cambr.) ; New Zealand
(Gray , Hutton).
Zooecia constituting erect cylindrical branches, arranged in 4, 6 or more series, and
opening all round the branch. In 4-serial specimens the arrangement is Farcimznaria-like., the
branch being a 4-sided prism, each surface formed by one of the rows of zooecia. Most of the
free surface of the zooecia constituted by a frontal membrane, into which the parietal muscles
are inserted; the frontal membrane not extending to the proximal end, which is the "peduncle"
of Hincks (1884). Orifice resembling that of Flustrella hispida, but without a clistinct upper
lip. The lower lip is well marked, and lateral lips are indicated. Spines absent.
The specimens described by Hincks were 6-serial. In colonies in the Cambridge Museum
which had formed part of Mr Hincks' collection I find this arrangement in most of the branches ;
but one branch is 4-serial. In a specimen from Port Jackson, in the same Museum, some of
the branches are also 6-serial ; but just before a bifurcation the branch may be widened and
flattened, as many as 5 rows appearing on each of the flat surfaces. But even in this colony
one of the branches is 4-serial. If thus appears that, although six rows are commonly present,
the number may be either greater or less. The Torres Straits specimen here described shows
an increase in the number of the rows to 8 in one place, just before a bifurcation.
The diameter of the branch drawn (fig. 2) is 600 u.. The middle zooecium is 1030 u>
long, and its frontal membrane is 370 u. broad at its widest part.
The present species has not been found in the 'Siboga' dredgings ; but I include it here
on the evidence of a single fragment obtained by Dr A. C. Haddon in Torres Straits. It belongs
to a series of species, of erect habit and with more or less cylindrical branches, which appear
to be characteristic of Southern waters. In addition to the species here described, this assemblage
39
40
includes Flustrella binder i HarveyM and F. hispida var. cylindrica Hincks 3). From these two
species8) E. blainvillii differs in the absence of spines.
Verrucularia dichotoma was originally found at Algoa Bay and was described as an
Alga bv v. Sihr: and there has been considerable doubt with regard to its systematic position.
Busk (1861) referred it and Flustrella binderij with some hesitation, to the Cheilostome genus
Farciminaria\ but later (1884) he accepted v. Suhr's genus, which he placed in the Farcimi-
nariidae. Hincks (1884) gave reasons, which he maintained in his later account (1893), for
considering V. dichotoma a species of Flustrella, in opposition to the opinion of Busk and
MacGillivray (1887, 1890). In placing it in the Flustrellidae I agree with Hincks ; and after
examining the species above mentioned, it seems to me that the structure of the orifice
corresponds closely with that of the common European F. hispida.
Goldstein *) has suggested that Farciminaria may be a synonym of Elzerina, but if
Lamouröux' species is the one here considered, this opinion cannot be maintained. In the original
account of E. blainvillii the "substance" of the colony is described as being "presque membra-
neuse"; — suggesting an absence of calcification. The specimens are said, moreover, to have
been found on sea-weeds from Timor and Australasia; and their occurrence in the Malay
Archipelago is a reason for referring the Torres Straits specimen to the same species.
It may be noted that Flustrella flabellaria Kirkpatrick '), from the China Sea, appears
to be a Pkerusa, so far as I can ascertain from a figure given by that author and from the
mounted slide (89.8.21. 128) in the British Museum.
A few words may be added with regard to the unfortunate results of attempting to
apply the Law of Priority strictly to a group like the Polyzoa, in which the earlier names
cannot always be identified with certainty. The familiar and universally employed generic name
Flustrella Gray, 1848, is antedated by Flustrella Ehrenberg ü), 1839. Ehrenberg's name was
introduced for a Cretaceous species, Flustrella concentrica, a member of a Family Asterodiscina,
where it is associated with Luuulites and Cupularia, two genera of Selenariiform Polyzoa, and
with Orbitulites, one of the Foraminifera. On p. 132 of Ehrenberg's work, E. concentrica is
defined as "microscopica cellularum minutissimarum laevium seriebus concentricis, interdum spira-
"libus, apertura singularum parva rotunda". In the 'Tabellarische Charakteristik d. Bryozoen-
Classe' following p. 120, the words "forma indeterminata plana (libera)" appear to indicate that
Flustrella was introduced for a Selenariiform Polyzoon ; but, so far as I am aware, Ehrenberg's
genus has not been recognised by later writers. Gray 's name has in fact held the field for 66
years, and the Family Flustrellidae, based on it, has also come into universal use. I venture
to think that this is a case in which the Law of Priority should be ignored 7).
1) Cf. Busk, 1861, Quart. J. Mier. Sci. (N. S.) I, p. 156.
2) Hincks, T., 1884, Ann. Mag. Nat. Mist. (5) XIII, p. 365: see P. H. MacGiu.ivkay, 1887, Trans. R. Soc. Vict., XXIII, p. 220.
3) Echintlla placoides Korotneff (Biolog. Centralbl. XXI, 1901, p. 311), a spinous form from Lake Baikal, may possibly belong
to the same assemblage. ANNANDALE ("Fauna Brit. Intl.", " 1'. W. Sponges . . . and Pol.", 191 1, pp. 199, 202) regards it, perhaps rightly,
synonym of Hislopia.
4) Goi.dstein, J. R. V., 1882, "New Species Bry. Marion Islands", Trans. R. Soc. Vict., XVIII, p. 44, note.
5) Kirkpatrick, R., 1890', p. 23, PI. IV, figs 3, 3«.
6) EHRENBERG, C. G., 1839, "Üb. d. Bildung d. Kreidefelsen", Phys. Abh. k. Akad. Wiss. Beilin, A. d. Jahre 1838, p. 132 and Tab. I.
7) Flustrella D'Orbigny, 1850 — 1852, p. 2S2, is a third introduction of this generic name.
40
4'
It seems to me further that it would be undesirable to attempt to replace Flustrella
Gray by either Elzerina Lamouroux, 1816, or Verrucularia v. Suhr, 1834. There is some
doubt as to the species which Lamouroux intended ; while it can hardly be considered proved
that the branching- Southern species which I have included under Elzerina really belong to
the same genus as the European species. Even if it be admitted that Elzerina should be
placed in the Flustrellidae, it may still be considered that the zoarial form justifies its separation
from Flustrella. To refer the branching species to the same genus as the encrusting European
species may be to suggest a wrong idea with regard to the geographical range of Flustrella.
F. kispida has a remarkable bivalved larval form which, so far as is known, is almost peculiar
to that species. In the absence of information with regard to the larvae of the Southern species,
it would be unsafe to assume that they are really congeneric with F. kispida. It thus appears
to me preferable to retain Gray's name for F. kispida- ; and in view of the fact that this has
been generally accepted for so many years I venture to think that it would be better not to
disturb it even if it were admitted that it is synonymous with Elzerina and Verrucularia.
Elzerina may perhaps be retained for the erect, branching Oriental and Southern species.
PALUDICELLEA , STOLONIFERA and VESICULARINA.
Before discussing the systematic arrangement of the Ctenostomata included below under
these groups, the method adopted for mounting preparations may be indicated.
In spite of the delicate nature of the body-wall in the genera here considered ; — which
might have been expected to render them easily permeable to reagents ; — it is well known
that the microscopical preparation of this group of Polyzoa is attended with special difficulties.
It is most desirable to find some method of mounting a type-specimen in Canada balsam,
and thus to give it as permanent a character as possible; but, in spite of many experiments
which I have made, I have been unable to discover any process by which the transference
into that substance can be effected satisfactorily. There is no difficulty in obtaining material
which is well preserved in either spirit or formalin. But on using oil of cloves, or creasote,
a shrivelling of the body-wall almost invariably takes place ; suggesting that the cuticle is
specially impermeable to anything in the nature of an oil. This property increases with age,
so that while the very young zooecia can sometimes be transferred to the oil without any
undue amount of shrinkage, the older zooecia and the older parts of the "stolon" commonly
collapse when treated with a reagent of this nature. In some cases, moderately successful
results can be obtained by dispensing with the use of an oil and transferring the preparation
directly from absolute alcohol to Canada balsam dissolved in absolute alcohol, as recommended
in my paper on the Morphology of the Cheilostomata l). But even when this procedure is
foliowed the results are often very disappointing. In a few cases I have had some success by
ij Harmer, S. F., 1902, Quart. J. Mier. Sci., XI.VI, p. 264.
SIBOGA-EXPEH1T1E XXVIIItf.
42
treating the specimen tor some hours with an extremely dilute solution of Eau de Javelle,
with the view of increasing the permeability of the cuticle : - - a method which has been
recommended for the preparation of Nematoda : but the reagent in question readily destroys
the interna! organs if used too strong, and it is very difHcult to regulate its action in such a
way as to obtain a successful result.
I have tlius been obliged to depend mainly on glycerine-preparations. The specimen is
transferred from 70 p.c. alcohol into a solution of glycerine in 70 or 90 p.c. alcohol which is
sufhciently dilute to cause no appreciable contraction of the body-wall of the spirit-material. The
specimen is then left exposed to the air, in an open watch-glass, for about two days, during
which the greater part of the spirit evaporates; and is finally mounted in some of the glycerine
which has thus concentrated itself to a strength which is in equilibrium with the air.
For mounting the slides I have made use of a method which has been elaborated by
Mr C. F. Rousselet, to whom I have to express my acknowledgments for information which
he has kindly given me. A "cavity-slide", or one in which a depression has been ground, is
used for this purpose ; and the first operation is to make an extremely narrow ring of gum-
dammar dissolved in benzol, immediately outside the depression in the slide. This process is
effected with the aid of a turn-table ; and the benzol evaporates so quickly that the slide is
ready for use as soon as the ring has been made. The object of this procedure is to raise the
cover-glass (a circular one, the diameter of which is slightly greater than the ring of gum-
dammar) above the level of the slide. After placing the specimen, in glycerine, in the cavity of
the slide and putting the cover-glass in place, the excess of glycerine is removed as completely
as possible by means of a fine pipette or filter-paper. If this operation has been successfullv
accomplished a circular space is left between the gum-dammar and the edge of the cover-
glass into which the cementing substance can be introduced. The escape of the glycerine
is thus prevented by a circular ring of cement between the outer margin of the cover-glass
and the slide and forms a more efficiënt closure than in the more usual method of mounting,
in which the cement merely passes from the slide over the edge of the cover-glass to its
upper surface.
The cements used have been, following Mr Rousselet's advice; - - first a ring of "picture-
copal" varnish, allowed to dry for 24 hours; and after that several successive rings, added at
intervals of a day, of a mixture of equal parts of picture-copal varnish and gold-size. For
thicker specimens a glass-cell has been used, cemented to an ordinary slide, instead of the
cavity-slide. In order to avoid the escape of the cement into the cavity enclosed by the cell,
the glass-ring should receive several successive rings of the copal-gold-size mixture, on its lower
edge, several days before the cell is required. The attachment of the cell is then readily effected
by placing it in position on the slide and hearing it over a flame. Before the glycerine is placed
in the cell the upper surface of the glass-cell should receive a ring of the cement which should
be left for some hours, long enough to allow it to become partially dry. The cover-glass should
be slightly larger than the diameter of the cell, so that a narrow margin, to which the cement
can adhere on the under side, is left projecting beyond the cell. Several successive rings of
cement are added, as in the case of the cavity-slides.
43
It should be added that Mr Rousselet prefers formalin to glycerine as a medium for
mounting; but I have myself used glycerine on the ground that there is less risk of drying up
if the cementing process has not been completely successful.
The arrangement of the genera of non-encrusting Ctenostomata is admittedly a matter
of great difficulty; and they have been groupecl in a considerable variety of ways by different
authors. Waters ]) has pointed out that the Division Stolonifera, as used by rnany authorities,
includes more than one type of structure, and that its type-genus, as originally introduced by
Ehlers, is Hypophorclla. He accordingly suggests the following classification : —
Vesicularina. "Usually 8 — 10 tentacles ; a gizzard; the zooecia grow direct from the stem and
"have no independent movement". Genera: — Zooóotryou, Bowcrbankia, Amathia,
Vesicttlaria, ? Avenella, Buskia (B. soda/is), Cryplopolyzoon.
Stolonifera. "There is a delicate creeping rhizome, which at intervals expands, and from
"these expansions the zooecia arise usually in pairs; no gizzard". Genera: —
Hypophorella, Farrclla, Valheria, Mimosella, Triticella, Cylindroecium, Buskia
(B. setigerd).
Waters is uncertain where to place Arachnidium (which may be allied to Cyliiidroeciitiu),
Victorella and Arachnoidea.
Rousselet 3) has proposed the association of the fresh-water genera Paludicella, Victo-
rella, Pottsiclla and Arachnoidea, with the marine genera Arachnidium and Cylindroecium 3),
under the name of "Cruciform Stolonifera"; for which the name Paliidicellea, already in use
with a more restricted meaning, might be employed. The study of the 'Siboga' material seems
to me to lend much support to this grouping, which is based on the fact that each zooecium
typically gives off a terminal daughter-zooecium, and another on each side, more or less at
right angles to the parent; thus producing a cruciform arrangement of uniserial zooecia. The
term "stolon", which is sometimes applied to the narrow tubes which connect the expanded
parts of the zooecia, is not very suitable in this connexion, since the tubes in question are
merely the proximal parts of the zooecia themselves.
Group B. Paludicellea.
Paliidicellea Allman, 1856, «Mon. F.-W. Pol." Ray Soc, pp. 10, 76.
Paludicellina Jullien, 1S88, "Miss. Sci. Cap Horn", VI, «Bry."', p. 7.
Paludicellina Annandale, 191 1, "Faun. Brit. IncL", "Freshwater Sponges . . . and Pol.", p. 190.
In this division may provisionally be associated the Families Paludicellidae, Victorellidae,
Arachnidiidae and Xolellidae (= Cylindroeciidae, auctt.), the first of which does not come within
the province of this Report.
1) Waters, A. W., 1910, pp. 23S, et seq.
2) Rousselet, C. F., 1907, Proc. Zool. Soc, p. 252.
3) Described as KoUlla in the present Report.
43
44
Fam. i. Victorellidae Hincks.
Homodiaetidae Saville Kent, 1S70, Quart. J. Mier. Sci. (N. S.) X, p. 35.
Victorellidae Hincks, 1880, "Hist. Brit. Mar. Pol.", p. 558.
Victorellides Loppens, 190S, "Bry. d'eau douce", Ann. Biol. lacustre, III, pp. 149, 170, 171.
Paludicellidae (part), Annandale, 191 1, t. cit., p. 191.
The present Family was originally defined, unsatisfactorily, by Saville Kent, under the
name Homodiaetidae, for Victorclla pavida, described in the same paper as a new genus and
species. Kent's Family-name was replaced by Hincks, in 1S80, on the ground that a Family-
designation should be taken from a typical genus. Annandale has recently proposed to include
the Victorellidae in Paludicellidae ; but this association has been criticised by Braem *), on the
grounds that Victorella has a typical development of the funicular system, and that this is not
the case in Paludicella ; and that the former produces larvae, while the latter lays eggs 2).
Loppens (1908) has called attention to resemblances between Victorellidae and Cylindroeciidae.
Victorella Saville Kent.
Victorclla Saville Kent, 1870, Quart. J. Mier. Sci. (N. S.) X, p. 34.
Victorclla Hincks, 1880, "Hist. Brit. Mar. Pol.", p. 559.
Victorella Kraepelin, 1887, Abh. Natunv. Ver. Hamburg, X, p. 93.
Victorella Rousselet, 1907, Proc. Zool. Soc, p. 255.
Victorella Braem, 1911, Trav. Soc. Imp. St Pétersbourg, XLII, Lief. 2, p. 30.
Victorella Annandale, 191 11, "Fauna Brit. Ind.", "Freshw. Sponges [etc.]", p. 194.
Victorella Annandale, 19112, "Syst. Notes Cten. Pol.", Ree. Ind. Mus., VI, p. 195.
For many years after its first description by Saville Kent the genus Victorclla was
only known to include the single species V. pavida, which has been recorded from brackish
water in England, Germany and Australia 3). In 1887, Kraepelin1) described a new species,
under the name of Paludicella mülleri, from the Ryckfluss at Greifswald, N. Germany, in a
locality where the percentage of salt in the water was about 0.3 p.c. (p. 95). This species has
been referred by later writers (Braem, p. 33, Annandale) to Victorella. In 1907 V. symbiotica
was described by Rousselet from Lake Tanganyika. In 1 9 1 1 Braem gave an account of another
new species, V. Continental is, from Lake Issyk-Kul, in Turkestan, from water which is described
(p. 4) as salt, but to so small an extent that cattle will drink it. The genus has also been
recorded from the salt lake Birket-el-Ourun in Egypt. Annandale 5) refers specimens from this
locality to V. symbiotica, while Braem (t. cit., p. 33) states that those he has examined belong
to the same group as V. pavida, etc, and that V. symbiotica belongs to another group,
characterised by a different structure, not particularised, of the alimentary canal. Annandale 6)
has described a new species, V. bengalensis, from fresh and brackish water pools in India.
1) BRAEM, F., 1914, "Die Knospung v. Paludicella", Avch. f. Hydrobiol., IX, p. 546. n.
2) Ibid., 1896, "Die geschl. Entw. v. Paludicella" ', Zoolog. Anzeiger, XIX, p. 55.
3) WHITELEGGE, T., 1890, "List Mar. and F.w. Inv. Fauna Port Jackson", J. Proc. Roy. Soc. N. S. Wales, XXIII, p. 322,
(Cook's River).
4) Kraepelin, K., t. cit., p. ióo, text-figs on p. 159.
5) Annandale, N., 191 i2, p. 197.
6) Ibid., 19111, p. 195; 191 12, P. 197.
44
45
It follows from the above summary that the genus Victorella has nearly always been
found in inland brackish waters, although Annandale l) states that it is known to exist in the
littoral zone of the sea in Europe. The occurrence of the following species among the 'Siboga'
dredgings is thus of special interest. I record it as : —
i. Victorella sibogac n. sp. (PI. III, figs 12 — 15).
Type. 45LB. Stat. 64. Kambaragi Bay, Tanah Djampeah, o — 32 Metres; coral, coral-sand ; trawl,
dredge and shore-exploration (On a chitinous tube, perhaps of a Phyllocliaetopterus).
331. B. Stat. 64. do., do. (on Tubucellaria fusiformis).
337. K. Stat. 71. Makassar, o — 32 Metres; mud, sand vvith mud; dredge and shore-exploration.
(Also on slide 337. G.1, Nolella papuensis, from the same Station). (On the axis of an
Alcyonarian).
The original label of 451. B. stated the depth as 32 Metres; and that of 331. B. as
30 Metres. It thus appears that the specimens were dredged and were not from the "shore-
exploration" included in the official list of the Stations. 337. K. was marked 27 — 36 Metres.
Zoarium partly adherent, partly giving rise to erect growths. Zooecia large, very variable
in form; sometimes with the erect portion less developed than the adherent part (fig. 13), but
in other conditions with the adherent part evanescent (fig. 1 2). Daughter-zooecia produced
sparingly, from the side of the parent, or as terminal buds continuing its direction. Distal end
of the zooecium more or less rounded, the orifice only slightly quadrangular. Polypides with
numerous tentacles.
Previous authors are agreed in stating that the genus Victorella is characterised by
having 8 tentacles. In the present species the number is certainly much larger. I have been
unable to ascertain it accurately, but I think it is safe to say that it is not less than 16, and
probably more than 20. I can find no other character which would justify the separation of
this species from Victorella, the generic diagnosis of which will accordingly have to be amended,
with regard to the number of tentacles, if V. sibogae is to be included in it. In certain other
respects, as in the mode of branching and in the characters of the oesophageal limb of the
alimentary canal, the present species shows very definite resemblances to others which are
referred to the same genus.
The cuticle of V. sibogae is thick and brownish in the older parts, but it is sufficiently
transparent to allow the internal structures to be seen fairly well in mounted specimens. The
condition is, however, very different from the "extreme transparency" alluded to by Hincks in
V. pavida. Numerous transverse or circular wrinkles run across the ectocyst (fig. 14), to a
degree which varies in different zooecia and in different parts of the same zooecium.
The general architecture of the colony conforms closely with the type prevalent in the
genus. The difference between attached and free parts is so great that it might be supposed
at first sight that two species were present; but this is quite in agreement with what is known
of other species. In the basal parts of the colony (fig. 13) a considerable proportion of the
1) Annandale, N., 191 i2, p. 197.
45
46
zooecium is adnate, the tree part bcing less than half the total length. In this condition, terminal
and lateral buds are developed; the former from the distal end of the attached part of a
zooecium, and the latter from one of its sides. A single zooecium may give off either one or
two lateral buds ; in the cases observed, from one side only. All the zooecia in fig. 1 3 are still
in the adnate condition. In other parts, however (fig. 12), the zooecia increase greatly in length,
and become completely erect As described by Annandale '), in his account of the generic
characters, the adnate part of the zooecium becomes hardly distinguishable, or quite indistin-
guishable, in this state. Lateral buds may be developed at either end of the zooecium (the basal
and distal buds of Annandale -) ), or from a more central position ; all these conditions being
shown in fig. 12.
The free portion of the zooecium is circular or oval in transverse section; and the orifice,
which is at the end of this part, is not sharply quadrangular, although showing indications of
having this shape (figs 14, 15). The free end of the zooecium appears rounded during retraction,
so long as the polypide is functional. When the polypide begins to degenerate, the end may
become pointed, as shown in one of the zooecia in fig. 12, where the degeneration is indicated
by the atrophy of the vestibule; the tentacles and alimentary canal not showing much change.
This difference of shape betvveen zooecia with and without a functional polypide is in accordance
with what is known of other Ctenostomes; and is alluded to below, under Valkcria atlantica.
The condition in which the zooecia have been preserved does not enable me to make any
statements about the collar.
The parietal muscles 3) are unusually strong, no doubt in correlation with the thickness
of the ectocyst. In the proximal part of the zooecium (fig. 15) their groups include a considerable
number of fibres, which are so long that in a side view of the zooecium they may appear like
bundies of circular muscles. The groups extend along the whole length of the zooecium, the
number of fibres becoming less at the oral end.
The polypide is relatively small, although sometimes considerably larger than in fig. 15, so
that the body-cavity is spacious. The vestibule is of moderate length, and the retracted tentacles
are bent at their distal end. Hincks (1880) has placed Victorella in his Campylonemidan Series,
characterised (p. 550) by having 8 tentacles, two of which are always everted, in the protruded
condition, and by having no gizzard. Kraepelin (1887, p. 95) has stated that the tentacles are
all straight, and are therefore not Campylonemidan, in Hincks' sense; while Loppens (1908, p. 9)
makes a similar statement. It is obvious that the large number of tentacles possessed by the
present species is a further departure from Hincks' definition. In correlation with the large
amount of space available in the body-cavity, the alimentary canal of the retracted polypide
retains its U-like character, instead of having the oesophagus bent on the pharynx. The statement
that there is no gizzard has been criticised by Annandale, who describes a part to which he
1) Annandale, X., 1911', p. 194.
2) Ibid., 19112, p. 195.
3) In his account of the genus in the "Fauna of British India", Annandale states (p. 195) that parietal muscles are absent.
It may be worth while to point out that an erratum-s\ip deletes this statement; slnce insertions of this kind are very apt to be lost or
overlooked. In his "Syst. Notes'', published in the same year, ANNANDALE has made use of the parietal muscles for the discrimination
of species of the genus..
46
47
gives that name !). As bearing on the correctness of the reference of the present species to
Victorella, the existence of a region (fig. 15) corresponding with Axxandale's "gizzard" may be
specially emphasised. I am not sure, however, that the part in question really corresponds with
the gizzard of other Ctenostomata, as it would appear to represent the oesophagus of other
forms; the preceding part of the alimentary canal, between it and the tentacles, corresponding
with the pharynx.
The existence of a distinct funiculus does not seem to be at all common in marine
Polyzoa, in most of which there is a more diffuse arrangement of the "funicular tissue". Victo-
rclla, however, is described as having a single funiculus, and it is therefore important to notice
that the present species forms no exception to this rule. In cases where I can obtain distinct
evidence, I find that the interzooecial septum is perforated by two pores, which lie side by side.
Through each of these passes a strong strand of the funicular tissue, the two cords uniting,
shortly after penetrating the septum, into a single cord which extends through the body-cavity
to be insertecl into the wall of the stomach.
Regeneration of the polypides is indicated by the fact that the zooeciurn often contains
one or two brown bodies (fig. 15).
The measurements of two of the zooecia figured, in y., are as follows, and may be taken
as a fair representation of the size :
fig. 15: length (not quite complete) 3100; breadth, 650.
fig. 14: length 1300; breadth, 600.
V. sibogae may be regarded as a specially robust member of its genus, as shown by the
large size of its zooecia, the thickness of its cuticle and the strong development of its parietal
muscles, and the large number of its tentacles. It appears to be nearly related to V. bengalensis
and V. continentalis, in both of which the number of tentacles is, however, stated to be 8; while
both are described as having winter-buds or resting buds. These have not been found in the
present species ; and it may be surmised from its occurrence in the sea that they do not occur in it.
It has often been assumed that Victorella is a form which has recently migrated into
fresh water, as indicated by its occurrence in brackish water. A different view is held by Braem,
who considers -) that its preference for brackish water, taken in conjunction with the localities
from which it has been recorded, indicates an ancient adaptation ; and that Victorella is in fact
a "relict" form which may still be found in localities which contain suitable brackish water.
The occurrence of a species of the genus in definitely marine localities appears to support the
older view rather than the one suggested by Braem ; unless indeed it were assumed that it
could have been derived from a form like V. bengalensis, living close to the sea and secondarilv
colonising it. Except on this view it would hardly be possible to assume, in the light of the
evidence afforded by V. sibogae, that most of the generic characters of Victorella had been
evolved in relation to its fresh-water habitat.
1) See especially Annandai.k, 1911', pp. 191, 195; where the "gizzard" 15 said not to function as that name would indicate,
but to be merely a chamber for the retention of solid particles: and 191 1-, PI. XIII, fig. 6, where the structure in question (6) is more
distinctly indicated.
2) t. cit., p. 34.
47
. '
48
One of the principal arguments relied on by Braem is the occurrence of V. continentalis
in Lake Issyk-Kul, a sheet of water which lies at a high level far in the interior of Asia. But
when it is remembered that I'. bengalensis gives rise to resting buds, it is not very unsafe
to suppose that these might be carried from one place to another, on the feet or beaks of
migratory birds, without losing their power of germination ; and that very long intervals might
be bridged over in this way.
Whatever view is adopted with regard to this point, it is interesting to find a marine
species of Victorella in a locality which may be regarded as central with reference to the Indian
Coast and Lake Tanganyika, two of the localities from which species of the genus have been
recorded. This fact becomes even more significant when taken in conjunction with the occurrence,
recorded below, in the Malay Archipelago of a species of Arachnoidea which appears to be
closely related to A. ray-lankesteri of Lake Tanganyika.
Fam. 2. Arachnidiidae Hincks.
Arachnidiidac Hincks, 1SS0, ''Hist. Brit. Mar. Pol.", p. 508.
Hislopiidae (pars), Annandale, 191 1, "Syst. Notes Cten. Pol.", Ree. Ind. Mus. VI, p. 197.
Zoarium adnate, branching typically in a cruciform manner, though frequently showing
some irregularity in this respect. Zooecia consisting of a narrow proximal part, often much
elongated, and of a much dilated distal part. Orifice subterminal, on a low papilla or a much
elongated peristome r). Genera, Arachnidium^ Arachnoidea.
Arachnidium Hincks.
Araclinidia Hincks, 1859, Rep. 28* Meeting Brit. Ass., Leeds, 1858, Notices and Abstr., p. 128.
Aracluüdia Hincks, 1859, Quart. J. Mier. Sci., VII, p. 131.
Araclinidia Hincks, 1862, Ann. Mag. Nat. Hist. (3) IX, p. 471.
Arachnidium Hincks, 1S77, Ann. Mag. Nat. Hist. (4) XX, p. 216.
Arachnidium Hincks, 1880, "Hist. Brit. Mar. Pol.", p. 508.
The first notice cited above mentions the description of "a new genene type, Araclinidia
" Hippothboides, a delicate Ctenostomatous Polyzoon, curiously resembling in general appearance
"the well-known Hippothod" . The second is a copy, not literally exact, of the first. A formal
diagnosis was published in the 1862 paper; but the information given in the 1859 notices is
probably sufficiënt to justify this year as the date of the introduction of the genus.
In 1877 Hincks uses the form Arachnidium, giving the derivation as "dim. of cï.puyyiov,
a spider's web", and stating that the name "Araclinidia" had been given "wrongly". I believe
that the amended form has been employed in practically all later accounts of the genus, and it
seems to me desirable to decide that the original name should be regarded as a lapsus ca/aun\
and that it may therefore be replaced by the correct form Arachnidium, in accordance with
1) I usc the term "peristome" as the equivalent of the free, terminal jiortion of the zooecium, hearing the orifice, without
implying that it is the morphological equivalent of the parts so designateil in Cyclostomata and Cheilostomata.
48
49
Art. tg of the Rules of Nomenclature 1). I must admit that this procedure involves stretching
the Rule, but it has the advantage of making the name of the genus next to be described
more different from that of the present genus than would otherwise be the case.
i. Arachnidium irregulare n. sp. (PI. III, figs 3 — 6).
376. B. Stat. 213. Saleyer, O — 36 Metres; coral-reefs, mud, and mud with sand. (On the dermal
membrane of a Siliceous Sponge).
Zooecia discrete, relatively large. Daughter-zooecia given off irregularly (hence the specific
name), separated from the parent by a diaphragm and at first narrow and cylindrical, then
expanding into a broad part, varying in form, and containing the viscera. Orifice borne on a
prominent, broad mammilla, situated subterminally, marked by concentric transverse lines, but
not angulated. Tentacles numerous.
The present species has been found once only. The spirit containing it had almost dried
up on arrival.
Hixcks (1862) stated, in characterising the genus, that "The Arachnidia may be
"regarded as an Alcyonidium with its cells detached from one another, and held together by a
"delicate thread". A irregulare conforms well with this definition: the "delicate thread" beinor
merely the proximal end of a zooecium. The length of this part is variable. The entire zoarium
is very closely adherent to the substratum, and the wide part of the zooecium is depressed,
ancl varies much in outline.
The cruciform disposition of the zooecia is usually apparent in Arachnidium ; and the
present species shows distinct traces of it (tig. 3), accompanied by some irregularity. Thus I
find that in A. irregulare there is commonly 110 symmetrically placed distal daughter-zooecium,
but that a zooecium is given off from each of the distal corners of the parent. This may be
the result of a displacement of the symmetry, by which one of the lateral zooecia appears to
form a pair with the distal zooecium (fig. 5). In other cases (fig. 4) two zooecia are given off
on one side and none on the other, the distal zooecium being median in the specimen figured.
In some cases (fig. 3) the two lateral zooecia are not opposite one another.
The orifice is borne on a massive mammilla, which is about half as broad as the attached
part of the zooecium, from which it rises nearly at right angles. The mammilla shows no tracé
of angulation, but is circular in transverse section, and is marked by conspicuous annulations.
The "collar" which is characteristic of the more typical Ctenostomata is not well showt) in the
preparations, but it appears to be present and of considerable size.
The tentacles are rather short and are numerous: — probably about 26 — 30. The
oesophagus is of moderate length (fig. 6) and the stomach and its caecum are large. In some
of the zooecia there appear to be several large embryos, one of which, contained in an ovisac
provided with radiating muscles, is shown in fig. 5. The vestibule has persisted, but the rest
of the polypide has degenerated.
In view of the variability of the proximal part of the zooecium it is not easy to state
teA
1) IXe Congres, Int. Zool., Monaco (1913), 1914, p. 899.
49
SIBOGA-EXPEDITIE XXVIII. /.
5°
the average length. The total length of the zooecium shown in fig. 5 is 1360 a. The length of
the narrow part of the proximal zooecium in tig. 4, measured to the level of the proximal lateral
diaphragm, is 600 u. ; and that of the broad part of the zooecium, from the same diaphragm,
is 1030 u.. The total length of the distal zooecium in the same figure is 1250 u,, while that
of the short zooecium shown in fig. 3 does not exceed 700 u., measured to the distal end of
the adhen-nt part. The zooecia are usually about 500 u in greatest breadth, and the diameter
of the narrow proximal portion is about 50 u.
A. fibrosum has been recorded by Miss Thornelv l), without description, from the Gulf
of Manaar.
Arachnoidea 2) J. E. S. Mooie.
Arachnoidia Moore, 1903, "The Tanganyika Problem", p. 297, text-fig. on p. 296.
Araclinoidia Rousselet, 1907, "Zool. Res. Third Tanganyika Exp.", Proc. Zool. Soc, p. 255,
PI. XIV, figs 5, 6.
Arachnoidea Annandale, 191 1, Ree. Ind. Mus., VI, p. 19S.
1. Arachnoidea protecta n. sp. (PI. III, figs 7 — 11).
378. E. Stat. 71. Makassar, o — 32 Metres ; mud, sand with mud, coral.
Zooecia discrete, relatively large, surrounded at their edges by irregular processes. Mode of
branching as in Arachnidium. Orifice borne on a long peristome, with thick chitinous walls, strongly
quadrangular when the polypide is retracted. Collar unusually large. Tentacles not less than 16.
The genus Arachnoidea was described by Moore (1903) from about 20 fathoms in Lake
Tanganyika, the single species found being designated A. ray-lankesteri. A later description
of the same species, also from Tanganyika material, has been given by Rousselet (1907).
Annandale (loc. cit.) has referred it to the Fam. Hislopiidae. LorPENS 3) considers Arachnoidea
inseparable from Arachnidium.
The occurrence of a marine species which can be referred to Moore's genus seems to
me of particular interest. But the resemblances between the •Siboga' specimen and those from
Lake Tanganyika are too striking to be overlooked ; and I feel no doubt that the forms from
the two localities must be placed in the same genus. As Moore pointed out, Arachnoidea closely
resembles Arachnidiuin in its general characters; but I think he was justified in separating it
from this genus by reason of the character of the peristome. In Arachnidium this part forms
a rounded mammilla, as shown in figs 3 — 6, referring to A. irregularc. Hincks accordingly
compared the genus with an Alcyonidium having discrete zooecia. In Arachnoidea, on the
contrary, the part in question is much elongated. lts wall is protected by a chitinous layer
which shows no tracé of the circular annulations in a relatively soft cuticle which were found
in the preceding species. In the retracted state, the peristome is quadrangular, the actual orifice
being perfectly square (fig. 11). It is true that in Arachnidium fibrosum Hincks4), the distal
1) Tiioknki.y, L. R., 1905, p. 127.
2) The name as originally introduced is Arachnoidia. I propöse to regard thi^ as a lapsus calami for Arachnoidea.
3) Loppens, K., 190S, "Bry d'Eau dovtce", Ann. Biol. lacustre, III, pp. 150, 154, 174.
4) Hincks, T., 1880, p. 511.
5°
5i
part of the zooecium is said to have a similar character; but specimens sent to the British
Museum (13.7. 10.4.) from Cullercoats, Northumberland, by Miss R. E. Roper1), and apparently
referable to A. jiörosum, have an orifice which seems to be of the typical Arachnidium form ;
the appearance of an elongated tube being obvious only in those individuals in which there is
some eversion of the kamptoderm.
The 'Siboga' specimens were growing on the surface of what seemed to be a fragment
of the axis of an Alcyonarian. I did not succeed in removing them sufficiently completely to
be able to describe the arrangement of the zooecia in any detail ; but there appears to be no
essential difference, in this respect, between the present species and Arachnidium irregulare.
In the existence of marginal crenulations, the present form closely resembles AracJinoidea ray-
lankesteri. The very large collar (figs 9, 10) distinguishes it from that species, and is the feature
referred to in the specific name suggested.
When fully protruded the collar forms an elongated cone, the sides of which are but
slightly divergent. It is supported by strong ribs, each of which ends distally in a strong tooth
(tig. 10), which is not quite terminal but projects into the cavity of the collar, the distal edge
of which is everted in such a way as to form a marked angle with the rest of the structure.
In zooecia in which the peristome is not so long (fig. 8) the proximal end of the retracted
collar may be seen projecting into the general zooecial cavity.
The specimens are not well enough preserved to allow me to describe the anatomy:
but I have evidence that the tentacles are numerous and not less than 1 6 in number.
The measurements (in u) of the specimens figured are as follows : - - total length of the
zooecium to the left of fig. 8, including its narrovv proximal part, 2700 ; without this part, 750 ;
breadth of the zooecium (fig. 11), 650 ; length of the peristome, when completely retracted,
400 — 700 ; diameter of the same part, 120; length of the collar, 370 ; diameter of the collar
at its base, j^\ at its distal end, 190; of the proximal part of the zooecia, 40 — 50.
A. protecta seems to be closely allied to A. ray-lankesteri. The marginal crenulations
of the zooecium are not quite so marked, or so regular as in that species, in which, moreover,
the peristome is less strongly chitinised and is not so quadrangular as in the 'Siboga' material.
Rousselet, in the paper cited above, figures no collar; and I have been unable to satisfy
myself of its existence in the specimens from Lake Tanganyika (Brit. Mus., 08. 5. 24. 9a, 9 b,
1 1 a — 1 1 c) described by that author. The differences in these parts seem to be a sufficiënt
justification for describing the 'Siboga' specimens as the representatives of a new species.
Annandale 2) has recently described a form to which he gave the name of Platypolyzoon
investigatoris n. g., n. sp., from the telson of Squilla investigatoris, obtained off the S. W. coast
of Arabia, in 110 fathoms; and considers that it is allied to Arachnidium or perhaps to Arach-
noidea. In the great length of its peristome it resembles the latter genus, from which it differs
in the characters of the attached part of the zooecium.
1) Roper, R. E., 1913, Dove Marine Laboratory, Cullercoats, Report for year ending June 1913, p. 53, PI. II, figs 2, la.
2) Annandale, X., 191 2, Ree. Ind. Mus. VII, Part if} N° 13, p. 124.
51
52
Fam. Nolellidae (new name).
Cylindroeciidae Hincks, 1880, aHist. J5rit. Mar. Pol.", p. 534.
If, as explained below, the familiar name Cylindroecium should be replaced by Nolella
Gosse, it appears to follow that the Family-name Cylindroeciidae, used by Hincks and other
authors, should be replaced by Nolellidae, taken from the type-genus.
The form which I describe below as Nolella annectens is so similar to Arachnoidea on
the one hand and to species of Nolella on the other hand that it is difficult to decide to which
of the two genera it should be referred. In its young state (PI. IV, hg. 1) this species may
show a strong resemblance to Paludicella. Somewhat later, with a greater development of the
peristome (hg. 8), the zooecium becomes very similar to Arachnoidea ; while in its final state
(fig. 3) it may be considered definitely a Nolella. In an intermediate condition of development
it diiïers from Arachnoidea mainly in being less dilated in its adnate portion.
Nolella annectens thus supports the view that the Families Paludicellidae, Arachnidiidae
and Nolellidae should be associated together as proposed by Rousselet l). Waters ~) is also
inclined to regard Arachnidium as allied to Nolella.
The resemblance of Victorella to Nolella [Cylindroecium) was remarked by Kraepelin3);
and this view has been supported by Delage and Hérouard 4) and by Loppens 5).
The Family Nolellidae is characterised by the great development of the peristomial part
of the zooecium. This region is typically much elongated and its ectocyst frequently includes
muddy particles. The adnate portion of the zooecium is represented by a delicate stolon-like
tube and by the base of the peristome, into which it usually passes abruptly, although it more
rarely dilates gradually as it approaches this part. The branching is of the cruciform type.
Gizzard absent. In some cases the embryos are known to pass separately into hernia-like
evaginations of the peristome.
The general structure of the zooecium in this Family is well shown in figures by Calvet i;).
Nolella P. H. Gosse.
Nolella Gosse, 1855, Ann. Mag. Nat. Hist. (2) XVI, p. 35.
Nolella Gosse, 1856, "Man. Mar. Zool.", II, p. 21.
Cylindroecium Hincks, 1880, "Hist. Brit. Mar. Pol.", p. 535.
On p. 540 of his "British Marine Polyzoa", Hincks mentions Nolella stipala Gosse as
a doubtful species, which is "Probably referable to Cylindroechiin" . In his original account (not
referred to by Hincks) Gosse gives the following diagnosis : —
1) ROUSSELET, C. F., 1907, "Zool. Res. Third TanganyiUa Exp.", Proc. Zool. Soc. p. 252.
2) Waters, A. W., 1910, p. 241.
3) KRAEPELIN, K., 1887, "Deutsch. Süsswass. Bry.'', Abh. Naturw. Ver. Hamb., X, p. 16S.
4) Delage, V. and Hérouard, E., 1897, „Traite Zool. Concrete'', V, p. 90.
5) LOPPENS, K., 1908, "Bry. d'eau douce", Ann. Biol. lacustre, III, p. 149.
6) Calvet, I.., 1900, "Contr. Hi>t. Nat. Bry. Eet. Mar.", Trav. Inst. Zool. Univ. Montpellier, Mém. 8, PI. VII, figs 12, 13;
PI. IX, figs 10, 11.
5-
''Genus Nolella (mihi).
"Cells erect, subcylindrical, springing singly, but closely, from an undefined polymorphous
"incrusting mat; tentacles eighteen, forming a bell. Name from nola, a little bell".
The zooecia of N. stipata are described as being about 1/36 inch long, whitish and
sub-opaque; the opacity making it impossible to ascertain whether a gizzard is present or not.
The number of tentacles is stated to distinguish it from all recognised genera except Avenella.
The emphasis laid by Gosse on the number of the tentacles, taken in conjunction with
his remarks on the opacity of the body-wall and with his figure (PI. IV, fig. 29) leave little
doubt in my own mind that the form described by him is identical with Cylindroecium giganten»/,
which Hixcks describes as possessing 18 — 20 tentacles. The conclusion that Cylindroecium must
be regarded as a synonym of Nolella thus appears to me in the highest degree probable.
Nolella stipata had previously been noticed by Bcsk1), with a reproduction of Gosse's
figure, in his original account of Farrella (Cylindroecium) gigantea.
Representatives of the present genus have been described from Oriental waters by
various authors ; and it is probable that some of these belong to one or other of the two
species here considered. In several of these cases 110 description or figure is published; and I
must content myself with giving the following references to the accounts in question, without
attempting to decide to what extent they correspond with the species described in this Report :
Cylindroecium papuense Kirkpatrick, 1888, "Pol. Port Phillip", Ann. Mag. Nat. Hist. (6) II,
p. 20 (Victoria).
Cylindroecium giganteum Hincks, 1887, "Pol. Hydr. Mergui Archipelago", J. Linn. Soc, Zool.,
XXI, p. 128 (said to be indistinguishable from British specimens).
Cylindroecium giganteum Waters, 1910, "Rep. Mar. Biol. Sudanese Red Sea" "Bry. II",
J. Linn. Soc, Zool., XXXI, p. 251 (diameter of stolon, 0.03 — 0.04 mm.).
Cylindroecium giganteum Thornely, 191 2, "Mar. Pol. Ind. Ocean", Trans. Linn. Soc. (2) Zool.,
XV, Pt 1, p. 157 (no description).
Cylindroecium giganteum Waters, 1914, ''Mar. Fauna Brit. E. Afr.", Proc. Zool. Soc, p. 854.
Cylindroecium dilatatum var., Kirkpatrick, 1890, "Hydr. Pol. China Sea", Ann. Mag. Nat. Hist.
(6) V, p. 17 (no description; the mounted specimen (Brit. Mus. 89.8.21.67.), from Tizard
reef, has deteriorated to such an extent that no conclusions can be drawn from it).
Cylindroecium dilatatum Thornely, 1905, HERDMAN's "Rep. Pearl Oyster Fish. G. of Manaar",
Publ. by the Roy. Soc, Suppl. Rep. XXVI, "Polyzoa", p. 128 (no description).
Cylindroecium dilatatum Thornely, 1907, "Rep. Mar. Pol. Ind. Mus.'". Ree Ind. Mus., I, Pt 3,
N° 13, p. 196 (spinous dilatations occur at the bases of the zooecia; — Mangalore, 26 — 31
fathoms).
Cylindroecium dilatatum Waters, 1910, t. cit., p. 251 (the zooecium has numerous fine latitu-
dinal lines; tentacles about 20 ; diameter of stolon about 0.02 mm.).
1. Nolella papnensis Busk. (PI. IV, figs 10 — 20).
Cylindroecium papuense Busk, 1886, Challenger Rep., Pt L, p. 38, PI. VIII, fig. 2.
2)*322. A. Stat. 50. Bay of Badjo, W. Coast of Flores, O — 40 Metres ; mud, sand and shells.
337. G. Stat. 71. Makassar, o — 32 Metres; mud, sand with mud, coral.
1) Busk. G., 1856, „Zoophytology", Quart. J. Mier. Sci., IV, p. 94, PI. V, fig. 4 (= X. stipata).
2) The specimens marked * «ere not examined as microscopical preparations.
53
54
56.1!. St.it. 8o. Borneo Bank, 2°25'S., ii7°43'E., 40—50 Metres; fine coral-sand.
'405. B. Stat. 91. Muaras Reef, E. coast of Borneo, 0—54 Metres; hard coral-sand, coral (on
a Sponge).
88. B. Stat. 115. E. side of Pajunga Island, Kwandang Bay; reef-exploration (on Porites).
108. P. Stat. 144. Anchorage N. of Salomakiëe (Damar) Island, 45 Metres; coral-bottom and
Lithothamnion (on a Hydroid).
•396. L. Stat. 162. Between Loslos and Broken Islands, \V. coast of Salawatti, 1S Metres;
coarse and fine sand with clay and shells.
130. H.\ Stat. 164. i°42.5 S., I30°47'.5E., 32 Metres; sand, small stones and shells. (130.0 is
1 30. O. ) on slide 1 30. M2, Bowerbankia ? imbricata).
*566. G. Stat. 204. Between Islands of Wowoni and Buton, 75—94 Metres; sand with dead shells.
376. A. Stat. 213. Saleyer Anchorage. o — 36 Metres; coral-reefs, mud and mud with sand.
(On slides 376. B.3, B.4, Araclinidium irregularé).
276. A.) Stat. 250. Anchorage off Kilsuin, W. coast of Kur Island, 20—45 Metres; coral and
*279. E. ) Lithothamnion.
"195. C. Stat. 274. 5°28.2S., i34°53'.9E., 57 Metres; sand and shells, stones.
Zooecia varying greatly in size, according to the state of their development, the ectocyst
usually striated transversely and including numerous particles of fine mud. Adnate portion
represented merely by the stolon-like proximal portion and by the base of the cylindrical portion,
which consists almost entirely of the peristome. This base gives off four or more stolon-like
threads, which may bifurcate after a time and ultimately pass into new peristomes. Fertile
zooecia containing several embryos near their distal end, and often very irregular in outline in
the region of the ovary or embryos.
The specific differences between the forms which have been referred to Nolella or
Cylindroecium are at present very obscure; particularly if I am right in believing that the same
species may be extremely variable, in respect of the size of the zooecia, according to its degree
of development. I have referred the present species to N. papuensis Busk mainly on account
of the identity in locality; although an examination of Busk's specimens in the British Museum
( s - . 12.9. 941. and 99.7. 1.4435.) leads me to think that the reference is correct. But the
question whether N. papuensis is synonymous with N. gigantca is so difficult that I am not
prepared to answer it. The fertile zooecia of the present species have a close resemblance to
those described by Prouho v), apparently from Banyuls, under the name of C. dilatatuiu. But,
from the examination of Hincks' type-specimen (Brit. Mus., 99. 5. 1. 1510), unfortunately a dry
specimen from which little can be made out, I am not confident that Prouho's reference was
correct. The 'Siboga' specimens also show considerable resemblance to N. gigantea Busk 3), the
type from Tenby (S.Wales), in the British Museum (99.7. 1.4434.) being much like the large
'Siboga' specimens. Busk does not mention the embryos, several of which are, however, present
in his specimen, although they appear to contain more yolk than in the 'Siboga' form. Another
slide (99.7. 1.4426) in the British Museum, also from Tenby, was labelled by Busk " Cylin-
droecium gigant cum var., seu C. dilatatum'\ The zooecia are not more than half the length
of those on the other Tenby slide; but this may merely indicate that the British species is as
variable in length as the Malay form.
1) Prouho, II., 1S92, Arch. Zool. Exp. (2) X, p. 626, PI. XXIV, figs 14—17.
2) Busk, G., 1856, »Zoophytology", Ouart. J. Mier. Sci., IV, p. 93, figs 1 — 3.
54
55
I have had great difficulty in deciding whether all the specimens indicated above belong
to a single species or not. Hut as I have been unable to find satisfactory distinguishing features,
it seems best to consider that only one species is represented; and that the differences observed
depend on the state of development.
This conclusion is supported by the fact that the zooecia on the same slide, and almost
certainly belonging to the same species, vary within wide limits. The young zooecia are relatively
short; while the old ones may be extremely long1). Some of the oldest zooecia, indicated as
such by the large amount of mud in their ectocyst, are, however, not longer than many of the
young individuals. The evidence obtained points to the conclusion that on the degeneration of
the polypide the zooecia may shorten considerably, partly by contraction and probably in part
by the loss of the distal end of the peristome. In some of these cases (fig. 10) regeneration
ot the zooecium has occurred, the younger part being sharply marked off from the older part,
not only by a distinct line but also by having an ectocyst with few foreign inclusions. It thus
appears that in some cases the zooecium, after reaching a considerable length, may become
shorter on losing its polypide, and may then grow longer once more when regeneration
takes place. In other cases there is no great shortening when the polypide degenerates.
But as this process involves the loss of the vestibule, a new vestibule must be regenerated;
and this is probably accompanied by an increase in length of the peristome. Embryos have
only been found in large zooecia; a fact which may indicate that several generations of
polypides have been present, the zooecium having increased in size with each such occur-
rence. The extent of the differences observed is brought out by the series of measurements
recorded below.
The peristomes are for the most part cylindrical along their whole course. The orifice
is usually square; but this is less obvious than in the following species (iV. annectens). The
vestibule is rather short. The collar is presumably present, but I have not obtained certain
evidence of its existence. If present it must be very delicate, and thus strikingly different from
that of N. annectens. The tentacles are about 18 in number, and are either quite straight
during retraction or are bent at their distal ends (fig. 19). The oesophagus is short, and is
succeeded by a distinct proventriculus. The caecum of the stomach is long. Strong parieto-vaginal
muscles are inserted into the vestibule; and parietal muscles are present, their fibres being
generally arranged two in a group.
Daughter-zooecia, commencing as stolon-like threads, are given off from the base of the
cylindrical part of the zooecium. Four are often present, arranged in the typical cruciform
manner. But a complication is introduced by the fact that the "stolons" may bifurcate. If the
division takes place at their point of origin, the zooecium may appear to give rise to a larger
number of "stolons", up to as many as eight. Each such tube is separated from the parent-
zooecium by a diaphragm, although the tube which represents the proximal end of the parent-
zooecium may show no diaphragm. The zooecium in such cases consists of a thread-like proximal
1) A similar variability has been noticed by SMITT (1865, "Om Hafs-Bry. utveckl. och Fettkr.", Üfv. K. Vet.-Ak. Förh., Arg. 22,
p. 30; and 1S66, »Krit. förteckn." II, Arg. 23, p. 525, PI. XIII, fig. 39), in the species of the same genus described by him as Farrtlla
and Vesicularia fusca.
55
56
portion, dilating into what has been described above as the adnate portion of the peristome,
which gives off other "stolons".
Some of the 'Siboga' specimens show embryos, in the situation which has been indicated
bv previous authors, and in particular by Prouho j). The embryos develop, according to this
author, in the body-cavity, at the distal end of a zooecium, three or four being present at one
time. When the larva has reached its full development, it pushes out the body-wall of the
zooecium and thus gives rise to a hernia-like protrusion, into which it passes. The outgrowth
eventually bursts, the larva escaping directly to the exterior. Small tubes projecting from the body-
wall may remain as indications of the spots where larvae have escaped in this way. Prouho's
account appears to be based on the actual observation of the escape of the living larvae.
Specimens which agree closely with Prouho's description are figured in this Report
(PI. IV, figs 10 — 12, 17). In fig. 10 an additional length of peristome has been added by
regeneration. At the distal end of the older part an egg is visible; while at the distal end of
the regenerated part there are two embryos. In fig. 1 2 three embryos occur in a similar position,
In fig. 17 two embryos are seen, near the middle of the length of the zooecium, each enclosed
in a thin hernia-like protrusion of the body-wall. It may be assumed, in view of the evidence
brought forward by Prouho, that these would shortly have escaped to the exterior. In fig. 1 1
there is again evidence of regeneration of the distal end of the zooecium. The younger part
contains an embryo distally ; while more proximally it shows irregularities in its outline, indicating
that embryos have escaped in this region. An egg is present in this part, so that the process
would probably have been repeated here.
Figs 19 and 20 possibly represent a different species. The vestibule contains several
rounded bodies which may be eggs. I cannot obtain certain evidence on this point; but if the
bodies in question are really of this nature, the specimen (108. P.) probably belongs to a distinct
species, characterised by producing eggs which pass into the vestibule, either undergoing their
development there or reaching the exterior and developing in the water outside, as in certain
other Polyzoa {JMembranipora pilosa, Alcyonidium albidum, HypopJiorclla expansa) which have
been described by Prouho 3). To this list of species, in which the egg is laid and undergoes its
development in the external water, may be added Pahidicella articu/ata (Braem, alluded to
above, p. 44) ; while Calvet 3) states that this is the case in Alcyonidium cellarioidcs, Pherusa
tubulosa and Flustrella hispida. It may, however, be remarked that the inclusion of Pherusa in
this list is directly opposed to the account which has been given by Prouho *), who states that
the embryos develop in the tentacle-sheath of an atrophied polypide ; and that the development
of the embryo of Flustrella hispida takes place inside the zooecium, as I have myself observed;
while Mrs Pace 5) has published an account of this species which shows that the position in
which the embryos develop is identical with that described by Prouho for Pherusa.
1 PROUHO, II., 1892, uContr. Hist. Bry.", Avch. Zool. Exp. (2) X, p. 626, PI. XXIV, figs 14 — 17.
2) Ibid., t. cit , p. 607, and elsewhere.
3) Cai.vet, L., 1900, "Contr. Hist. Nat. Bry. Eet. Mar.", Trav. Inst. Zool. Univ. Montpellier, N. S., Mem. 8, p. 260.
4) Prouho, H., t. cit., p. 565.
5) Pace, R. M., 1906. "Early stages Dev. Flustrella hispidd\ Quart. J. Mier. Sci., I., p. 451.
56
57
The following measurements, in y., will give some idea of the great differences in the
size of the zooecia, in different conditions of their growth :
A. Length of young zooecium.
337. G. 276. A.
B. Length of old zooecium, with functional polypide.
C. Length of old zooecium, with degenerated polypide.
800
1520
2590
2800
130. H.
900
1520
108. P.
3280
3680
3920
. IOIO
D. Length of old zooecium, containing embryos.
3135
3360
E. Greatest diameter of young zooecium (A).
370
1 120
I055
I330
190 175
!/D
'/b
F.
Greatest diameter of old zooecium (B).
175
160
25 5
24O
29O
G.
Greatest diameter of old zooecium (C).
225
240
145
190
255
H.
Greatest diameter of old zooecium (D).
320
335
The measurements given under E — H belong to the same zooecia as those recorded
in A— D.
The longest zooecium measured (130. H.) is nearly 4 mm. long1). An old zooecium
which has lost its polypide, and probably the distal end of its peristome, from the same Station,
is no more than 370 u, in length; which is less than that of the young zooecia. The specimens
with embryos (337. G.) are both more than 3 mm. long. 108. P., which consists entirely of small
zooecia, gives measurements which fall well within the limits of the young zooecia in 130. H.
The greatest diameters recorded are from zooecia with embryos (337. G.).
2. Nolclla annectens n. sp. (PI. IV, figs 1 — 9).
380.I. Stat. 47. Bay of Bima, 55 Metres; mud with patches of fine coral-sand.
Zooecia extremely variable in form; with or without a proximal stolon-like portion. The
basal adnate portion is very variable in size and shape, and the free tubular part (peristome)
is equally variable in length and in the angle it makes with the adnate portion. The peristome
1) Cylindroecium allum^ desenbed by KlRKPATRICK (iSSS, Arm. Mag. Nat. Hist. (6) II, p. 19) from Port Phillip, Victoria, reaches
a length of 4.6 mm.
57
SIEOGA-EXrEDITIE XXVIII a.
5»
becomes quadrangular distally, with a thick cuticle, and the orifice is square. Collar very strong.
Three buds are given off typically from each zooecium ; ■ — one from the distal end of the
adnate part, and one from each side in the same region (exceptionally two on one side).
The specimens here described were deeply embedded in the substance of a Monaxonid
Sponge, which had to be dissected away in order to exposé the Polyzoon. The variable character
of the peristomial region was probably the result of the conditions under which the Polyzoon
was growing; since the length and direction of the peristome would no doubt be dependent
on the extent to which the zooecium was immersed in the Sponge.
In the youngest specimen drawn (fig. 2), the basal dilatation is represented by a triangular
swelling, in which a young polypide-bud is present. The terminal bud is already cut off by a
septum from the parent-zooecium. The lateral bud of one side is represented by a large pointed
process in which no septum has yet appeared ; while that of the other side is a smaller process
of the same kind which has not developed symmetrically with the other one. The peristome
has not yet made its appearance.
In a later stage (fig. 1) the zooecium is Paludicc/la-\'\ke in shape; being represented
almost entirely by a pear-shaped portion which is continuous proximally with a very long
tube, of stolon-like form. The tube is, however, not a stolon in the proper sense, since it
gives off no buds; but is to be regarded, as in other similar cases, merely as the narrow
proximal end of the zooecium. The peristome is a short rounded elevation, much like the
corresponding part of a Paludicella-zoozcmm. This specimen shows a terminal bud, but only
one lateral bud.
Fig. 8 represents a fully developed zooecium, in which the peristome is still very short,
but has the quadrangular form which is characteristic of the fully developed state. It will be
observed that this individual has no narrow proximal portion. The terminal and lateral buds
have been developed normally ; and it will be seen that the lateral buds originate in the
region of the distal part of the adnate portion ; an arrangement which may be noticed in other
dravvings. In some cases, however, as in fig. 3, the adnate part is hardly developed, and the
lateral zooecia then appear to be given off from the base of a long tubular, erect zooecium,
as in more typical species of Nolella. The varying length of the part which ma)- be considered
adnate is responsible for the varying position of the lateral buds, which come off from the
distal part of this region.
Figs 6, 7, 4, 5 show various conditions of the peristome, intermediate between figs 8
and 3. The peristome may either come off at a distinct angle from the adnate part (fig. 7)
or may prolong its direction, with only a slight curve (fig. 5). In fig. 4 the zooecium is pear-
shaped ; the broader part being represented by the adnate portion of the zooecium and the
narrower end by the peristome. In all these cases, the quadrangular shape of the distal
part of the peristome is well marked, and is associated with a thickening of the cuticle in
this region.
With the degeneration of the polypide further changes may take place. The quadrangular
form of the peristome is lost and the kamptoderm becomes everted as far as the base of
the collar, which projects freely to the exterior, though with some appearance of undergoing
5S
59
deo-eneration (fig. 9). The distal end of the zooecium is now rounded. In others the collar has
been lost, and has probably been thrown off1), all traces of the vestibule having also disappeared.
Some of these old zooecia are considerably longer than the individual shown in hg. 3, and a
polypide-bud, or a young polypide, may be visible in their interior. It is natural to suppose
that this would lead, in course of time, to the reconstitution of a normal orifice ; and some of
the variability in the length of the peristome may be the result of the successive formation of
one or two polypides: each such formation being accompanied by au increase in length of
the zooecium.
Fig. 4 shows the general structure of a zooecium. The strong collar is well seen in
its retracted position, and the vestibule is provided with very strong parieto-vaginal muscles,
running obliquely from the body-wall into the vestibule. The tentacles are moderately long;
and they appear to be not less than 16 in number. The examination of other specimens
shows that they are not less than 20, in some cases at least. The pharynx is short, and, in
its retracted condition, lies close to the proximal end of the zooecium. The oesophagus is of
oreat length and passes distally, to about the region of the middle of the bundie of tentacles.
Here it opens into a large proventriculus, which may represent the gizzard of some other
Ctenostomes. The proportions of this limb of the alimentary canal are different from those
shown by Calvet -), in a longitudinal section of N. dilatata. The stomach and its caecum are
large, and the rectum is slender. A series of strong parietal muscles is present, on each side,
in the proximal end of the zooecium. The examination of other specimens shows that the fibres
are arranged in groups.
The arrangement of the alimentary canal shown in fig. 4 is, however, not constant; and
in fig. 7 it may be seen that the apex of the retracted caecum is directed distally. In some of
the very long zooecia above described the alimentary canal has a U-like form, in correlation with
the large amount of space available. The great elongation of the oesophagus is a characteristic
feature of the present species.
The variability in the size and relative proportions of the parts of the zooecia will be
brought out by thé following measurements, in u. :
Fig. 1 ; total length of the zooecium, including its narrow proximal part, measured along the
curve, 4000 ; greatest diameter, 240 ; diameter of the narrow proximal portion, 40 — 60.
Fig. 3; total length, 2600 ; diameter of the distal half of the peristome, 150.
Fig. 4; total length, 1670; greatest diameter, 350; diameter of the distal end, 120.
(Not figured); total length of an old zooecium which has no narrow proximal part, and in
which the polypide has degenerated, 5920.
Length of peristomes, measured from the angle formed with the terminal daughter-zooecium : —
Fig. 1, 180; Fig. 8, 220; Fig. 7, 570; Fig. 4, 1000 ; Fig. 5, 1050 ; Fig. 3, 2150.
In several of its characters the present species shows a considerable resemblance to
1) Braem (1914, "Knosp. v. Pahtdictlla", Avch. f. Hydrobiol. IX. pp. 539, 540) has described the throwing off of the tentacular
apparatus, or a part of it, on the degeneration of the polypide in Pahidkdla.
2) Calvet, L., 1900, t.cit., PI. VII. tig. 12.
59
6o
Arachnoidea protecta ; and this is specially obvious in an intermediate state of development.
When the zooecia have reached their highest development they are hardly distinguishable from
a typical Nolella. In the yoting condition the zooecium may be Paludicella-Wke.
Group C. Vesicularina Johnston.
Vesicularina (pars), Johnston, 1847, "Hist. Brit. Zooph.", Ed. 2, p. 367.
Vesicularina Waters, 1910, "Rep. Mar. Biol. Sud. Red Sea", "Bry. II", J. Linn. Soc, Zool.,
XXXI, p. 240.
The genera included by Johnston in the Vesicularina were Serialaria {Amathia), Vesiai-
laria, Beania, Valkeria, Bowerbankia and Farrella. Of these Beania is now placed among
the Cheilostomata, while Valkeria and Farrella have been transferred by Waters to the next
division of the Ctenostomata. With these removals the Group remains almost identical with
Waters's use of the name.
The Vesicularina may be defined as consisting of those Ctenostomes in which there is
a relatively thick, branching, tubular axis, to which the zooecia 'are directly attached, by a
contracted, or an expanded base. The axis is usually divided by diaphragms into internodes,
each of which gives off a number of zooecia. Budding only from the axis. The number of
tentacles is commonly 8 — 10. Gizzard present in most of the genera, and perhaps in all.
Family Vesiculariidae Johnston.
Vesiculariadae (pars), Johnston, 1838, "Hist. Brit. Zooph.", Ed. I, p. 247.
Vesiculariadae (pars), Johnston, 1S47, Do, Ed. 2, p. 367.
Vesiculariadae (pars), Gray, 1848, "List. Brit. An. Brit. Mus.", I, pp. 94, 145.
Vesiculariidae Hincks, 1SS0, "Hist. Brit. Mar. Pol.", p. 512.
The Family, as originally introduced by Johnston, included Vesicularia, Serialaria,
Valkeria and Bozoeröatikia. In the second Edition of his work the list of genera is identical with
that given above for the Vesicularina. Hincks included Avcnclla in it; — an association on
which I am not able to express a positive opinion. Zoobotryon doubtless belongs to this family.
As defined by Hincks, the Family consists of Ctenostomes having "Zooecia contracted
"below, not closely united to the stem at the base, deciduous, destitute of a membranous area.
Zoarium repent or erect". Part of this definition must certainly be excluded, since the zooecia
in Amathia are not contracted proximally nor individually deciduous. The definition of the
Family may provisionally be given as identical with that of the Vesicularina, since it is at
present uncertain whether any of the genera referable to the group deserve separate Family
rank. Farrella, which may belong to this Family, is believed to have no gizzard, a structure
which is known to occur in most of the other genera.
In the group Paludicellea the appearance of a "stolon", in certain genera, is due to the
fact that the proximal ends of the zooecia are much narrower than the expanded distal ends.
In the Vesiculariidae nothine of the kind occurs. The axis is sharply marked off from the
6i
zooecia to which it gives rise ; and even if its internodes are to be regarded as specially modified
zooecia — a view which has been supported by some authors — there is no very direct evidence
that this is the case, beyond the fact that the body-wall and the body-cavity of the axis-
internodes do not difter in structure from the corresponding parts in the zooecia. It appears
to me that the natural view to take of these internodes is that they represent parts of the
colony which are not divided into zooecia, rather than that each of them is to be regarded as
the morphological equivalent of a single zooecium. It is characteristic of the Vesiculariidae that
each internode gives rise to a number of zooecia, the arrangement of which is distinctive of the
several eenera.
ö
Vesicularia J. V. Thompson.
Vesicularia J. V. Thompson (pars), 1830, "Zool. Res.", N° IV, Mem. V, p. 97.
Vesicularia Farre, 1837, Phil. Trans., p. 402.
Vesicularia Hincks, 1880, "Hist. Brit. Mar. Pol.", p. 512.
The genus Vesicularia was introduced by Thompson (1830), who included in it V. cuscuta,
V. spinosa, V. puslulosa and V. imbricata. Of these, the first was referable to Fleming's
earlier genus \ralkeria. Farre, in 1837, pointed out that V. spinosa differs from V. cuscuta in
possessing a gizzard; and Sertularia spinosa Linn. has accordingly become the genotype of
Vesicularia. The other two species included in the genus by Thompson are both referable
to Bowerbankia.
Vesicularia, as now understood, (cf. Hincks, 1880) includes species with ovate, deciduous,
distant zooecia, contracted at the base, the polypides possessing a gizzard. In this restricted
form it appears to include but few species.
Busk added two species in the 'Challenger' Report: — V. papuensis from 28 fathoms,
off the S. coast of New Guinea, and V. trichotoma from Bass Straits. The former at least
is included in the 'Siboga' Collection.
&■
1. Vesicularia papuensis Busk. (PI. VI, figs 1 — 4).
Vesicularia papuensis Busk, 1S86, Challenger Rep., Pt L, p. 36, PI. VIII, figs 1 — \c.
Vesicularia trichotoma Busk, Do, p. t>7> PI- VIII, figs 4 — 4.0.
307. A. Stat. 273. Anchorage off Pulu Jedan, E. coast of Aru Islands, 13 Metres; sand and shells.
Zoarium reaching a length of 60 mm. Stem attached by a tuft of delicate rootlets, which
become very fine by subdivision. Main stem covered by a thick cuticle, angulated at the nodes,
the angulation being alternately on opposite sides. Each internode gives off two ordinary branches
and sometimes a third which forms rootlets only. Stem jointed immediately on the distal side of
the node; a similar joint occurring at the base of each branch. The branches subdivide somewhat
differently from the main stem, as the principal axis is not prolonged, and the branching thus
appears to be simply dichotomous. Zooecia very readily lost, arranged in two alternating series,
on one aspect of the branch. Polypide possessing a gizzard and 8 tentacles. Vestibule and
collar rather short.
61
62
In the main stem the two branches are both given off on the side towards which the
angle of the stem is dirccted. The third branch, which gives rise to rootlets, may come off
opposite the two ordinary branches (and therefore from the re-entrant angle of the stem), or
quite asymmetrically. At the division of the branches the main stem is suppressed at each node,
so that the two branches of the new order are alone left. A branch may bifurcate successively
3 — 5 times. The main stem is traversed by a diaphragm, possessing a central perforation,
immediately beyond the node; and a joint is formed in this position by a diminution of the
thickness of the cuticle (fig. i). A similar development of a diaphragm and of a joint is seen
at the origin of each branch ; and the rootlets are also provided with a diaphragm at their
points of bifurcation. The main stem usually has no zooecia, though a few may occur; but the
presence of scars, sometimes in the form of small papillae (fig. i), indicates that zooecia have
been borne, in those positions, at an earlier period in the life of the colony.
The branches, on the contrary, are provided on one of their aspects with a doublé
series of zooecia, sometimes represented merely by scars (fig. i), which have a definite alternate
arrangement. A single internode usually bears from 3 to 5 pairs of zooecia, although the
number of zooecia is sometimes odd.
The zooecia are constricted at their base, as in other species of the genus, and are
thus readily deciduous. I have not found any evidence that a new zooecium can be developed
from an old scar :).
In form the zooecia are sub-cylindrical, the broadest point being in their proximal half
(figs 2, 4). They agree precisely with those shown by Busk in his fig. 1 b (PI. VIII) except that
they become quadrangular towards the orifice; which, in the retracted condition, is quite square.
This condition is not alluded to in Busk's description. The tentacles are 8 in number. The
pharynx is rather large and is succeeded by a narrow, elongated oesophagus, which (in the
retracted state) passes distally and then opens into a distinct gizzard, stated by Busk to be
absent. The stomach and its caecum are large. Parietal muscles are present in several groups,
usually of two fibres each.
The following measurements, in ij., of the several parts of the colony agree closely with
those recorded by Busk :
Length of an internode of the main stem, 1900; diameter, 250.
Length of an internode of a branch, . . . 1100; diameter, 100 — 180.
Length of a zooecium, 480 ; greatest diameter, 150.
A specimen from Japan, in the Collection of the University Museum of Zoology at
Cambridge -), is certainly referable to the present genus, but it appears to belong to a distinct
species, so far as can be ascertained from the mounted slide, which is not a very successful
preparation. The zooecia are distant from one another, the interspaces being nearly equal to two
1) LEVINSEN ("Régén. totale des Biy.", Buil. Acad. Roy. Sci. Danemark, Ann. 1907, Nu 4, p. 152) has observed this regeneration
in Valkeria and Bowerbankia.
2) Off Tokyo, 53 fathoms, A. Owston Goll., Reg. June 23, 1902.
62
63
diameters of a zooecium. They thus appear to be arranged in a single row along the branch,
and the alternate disposition is not obvious. The ends of the branches are rounded or truncate.
V. spinosa, the type-species, is distinguished by having " spinous" ends to the branches,
composed of small barren internodes which are acutely pointed. The zooecia are not obviously
paired, but occur in what looks like a single row, three being commonly developed by each
internode, at considerable distances apart. In this respect, and in the apparently uniserial
arrangement, it agrees with the Japanese specimen.
In two points — the character of the branching and the existence of a gizzard — the
account given above differs from Busk's description. But I have convinced myself, by a comparison
with the original material, that the 'Siboga' specimens, which are from a locality very near
that from which V. papuensis was first described, really belong to Busk's species-, and indeed
that they agree extremely closely with his specimens. The type-material, in the British Museum,
consists of a specimen, Sy. 12. 9. 938, in spirit; and a slide, 87. 12. 9. 937, which was no doubt
prepared from the spirit-specimen. In describing V. trichotoma, which immediately succeeds
V. papuensis in the 'Challenger' Report (p. 37), Busk emphasises the trichotomous division of
the primary branches as a peculiarity distinctive of the former species; and therefore as a
difference between it and V. papuensis, in which the zoarium is said to branch dichotomously
in one plane. It may be inferred with some confidence that Busk wrongly assumed the mode
of branching visible in the mounted slide of V. papuensis (which shows only a lateral branch
of the colony) to be the only mode occurring in the colony. But a reference to the spirit-
specimen shows that in the main stem the branching of V. papuensis is as trichotomous as in
the 'Siboga' specimen. The stem is in fact angulated at the nodes, from each of which a pair
of divergent branches, not in one plane, are given off. In some of the nodes the stem might
appear to dichotomise, if examined superficially ; but on closer inspection it is seen that in these
cases one of the two "branches" has a thick cuticle and in other respects shows itself to be the
prolongation of the stem. The other branch has the thinner cuticle and the simple dichotomous
mode of dividing which characterise the lateral branches in general.
With regard to the second point, the mounted slide, on which Busk probably relied,
does not show the gizzard convincingly. But the zooecia of the spirit-specimen are suffkiently
transparent to allow the gizzard to be seen quite distinctly. It is thus seen that V. papuensis
conforms to the generic diagnosis given by most authors, in possessing a gizzard.
V. trichotoma is represented in the British Museum Collection by three slides (87. 12. 9.
939; 99- 7- ï-430?; 99- 7- 1 • 43 10) which have unfortunately suffered greatly by the drying up
of the medium in which the specimens were mounted. The trichotomous branching described
by Busk is well seen, however, in those parts of the colony in which the stem is covered by
a thick chitinous ectocyst. The terminal branches are "furcate", as Busk stated; and there is
indeed no obvious difference between these forms and V. papuensis. The greater diameter of
the sterns mentioned by Busk as a distinctive feature of V. trichotoma seems to me also without
rauch significance; since, as in the type of V. papuensis, the diameter becomes less in passing
from the proximal to the distal parts of the colony. The balance of evidence appears to me in
favour of regarding V. trichotoma as a synonym of V. papuensis.
63
64
Amathia Lamx.
Amathia Lamouroux, 1812 '), "Extr. mém. class. Pol. Cor. non entièrement pierreux", Nouv.
Buil. Sci. Soc. Philomat. Paris, III, p. 184.
Amathia Lamouroux, 18 16, "Hist. Pol. Cor. Flex.", p. 157.
Amathia Lamouroux, 1821, "Exp. Méth.", p. 10.
Amathia Lamouroux, 1824, "Encycl. Méthod.", "Zooph.", p. 42.
Amathia Waters, 1910, "Rep. Mar. Biol. Sud. Red Sea", "Bry. II", J. Linn. Soc, Zool.
XXXI, p. 242.
Serialaria Lamarck, 1S16, "Hist. Nat. An. sans Vert.", II, p. 129.
The genus Amathia has formed the subject of special Memoirs by Tenison-Woods ")
and MacGillivray 3) ; the former author giving a valuable account of its history, and the latter
describing a number of Australian species. It is well represented on the Australian coast, but
the 'Siboga' collection includes only two species.
1. Amathia convoluta Lamx. (PI. V, figs. 1 — 5).
Amathia convoluta Lamouroux, 1S16, t. cit., p. 160.
Amathia convoluta Lamouroux, 1824, t. cit., p. 44.
Amathia convoluta Tenisoon-Woods, 1880, t. cit., p. 100.
Amathia convoluta MacGillivray, 1895, t. cit., p. 133, PI. A, figs 3 — $b.
Serialaria crispa Lamarck, 1816, t. cit., p. 131 (fide LAMOUROUX, 1824, who gives S. convo-
luta Lamk ,as a synonym of A. spiralis Lamx).
Amathia crispa Pergens, 1887, "Contr. Hist. Bry. et Hydr. récents", Ann. Soc. Roy. Malacol.
Belgique, XXII, Buil. des Séances, p. xc (= Sirinx circumplicata Desmarest and Lesueur,
p. LXXXVIII).
Amathia spiralis Busk, 1886 (nee Lamx), Challenger Rep., Pt L, p. 34, PI. VI, figs 2— 2b.
Amathia semispiralis Busk, 1886, t. cit., p. 56, PI. VIII, figs 3 — 3b.
Amathia semispiralis} Waters, 1S87, "Bry. N. S. Wales", Pt III, Ann. Mag. Nat. Hist. (5),
XX, p. 264.
? Amathia tortuosa Busk, 1886 (nee Tenison-Woods), t. cit., p. 34, PI. VI, fi^s \ — \b (regarded
by Waters, 1910, t. cit., p. 242, as identical with A. convoluta).
Amathia tortuosa Kirkpatrick, 1890, "Hydr. Pol. Torres Straits", Proc. R. Dublin Soc. (N. S.),
VI, p. 612.
The following records may belong to this species; but the amount of information given
is insufficiënt to enable any certain statement to be made : —
Amathia convoluta Hasvvell, 1S81, "Pol. Queensland Coast", Proc. Linn. Soc. N. S. Wales,
V, p. 43-
Amathia convoluta Whitelegge, 1889 [1890], "List Mar. & Fr.-water Inv. Fauna Port Jackson",
J. Proc. R. Soc. N. S. Wales, XXIII, Pt II, p. 292.
1) Miss Jelly (1889, p. 279) and NlCKLES and Bassler (1900, p. 510), in their bibliographical lists, mention the title of a
paper, without further reference, supposed to have been published by Lamouroux in 1910; and Tenison-Woods (1880, p. 97) gives the
title in more detail. This paper appears not to exist. It is true that Lamouroux himself (1824, p. 43) states that he "proposed" the genus
Amathia in 1910: but in his "Histoire des Pol. Cor. Flex.", 1S16, p. 187, he gives 1812, with the reference cited above, as the date of
the introduction of that genus; and in the Preface (p. vi) of the same work he explains that his memoir was presented to the French
Academy in 1S10, but actually appeared two years later (1812).
2) TENISON-WoODS, J. E., 1S80, "On the Genus Amathia", Trans. Proc. R. Soc. Vict., XVI, p. 97.
3) MacGillivray, P. H., 1895, "0n tne Australian Species of Amathia", Proc. R. Soc. Vict. (N. S.) VII, p. 131.
64
65
'I Stat. 27;. Oft" Pulu Jedan, E. coast of Aru Islands, 13 Metres; sand and shells.
544. A. ) / ° J J
^ '[ Stat. 274. 5°28'.2S., I34°53'.9E., 57 Metres; sand and shells, stones.
126. A.)
} Stat. 164. i°42.sS., I30°4/.5E., 32 Metres; sand, small stones and shells.
(sp. ?) 67. C. Stat. 99. Anchorage oft" N. Ubian, 16 — 23 Metres; Lithothamnion-bottom.
also the following specimens, in the Mus. Zool., Cambridge : —
'°°' | Torres Straits, A. C. Haddon ColL Reg. Feb. 24, 1898.
T. S. 100.
T. S. 2c
Queensland, Holborn Island, Port Denison. From the remains of W. A. Haswell's Coll.,
pres. by the Australian Mus., Reg. Oct. 23, 1899.
Zoarium large, erect, well branched, reaching a length of at least 132 mm. Stem thick
at its base, where it is strengthened by rootlets given off at various points, the vvhole comrflex
here reaching a thickness of 10 mm. The ordinary internodes, without the addition of rootlets,
measure about 400 — 550 u. in diameter. Zooecia biserial, arranged in an open spiral curving
round the stem, a considerable part of which is exposed. Each group of zooecia forms, as a rule,
from i1/.-, to 2 complete turns of the spiral, seldom as many as 2l/.2 turns. A thick diaphragm
typically occurs at the distal end of each group of zooecia, a short interval separating the group
from its successor. In branches which have reached the limit of their growth a spiral series
may be broken up into several separate groups, each of a few pairs of zooecia, and isolated
from one another by a considerable interval of bare stem. A branch is given off, on one side
only, immediately on the proximal side of a stem-diaphragm ; successive branches being alternate
and on opposite sides of the stem. The zooecia are completely connate along their whole length
when the kamptoderm is fully invaginated, and the edge of the entire series then forms an
angular groove, the deepest part of which corresponds with the line separating the two series-
of zooecia, the outer walls of which project so as to form two sharp ridges bounding the
groove. The oral wall of the zooecium thus slopes downwards into the groove; and the orifice
is situated on the distal part of the flat surface so constituted. In transverse section the
zooecium is much broader than long, and the basal attachment to the stem is thus oblong,
the two longer sides being the proximal and distal ones, the two series usually interdigitating.
Polypides with 8 tentacles and a gizzard.
The present species, which has a considerable resemblance to the Mediterranean A. semi-
convoluta Lamx, as figured by Heller x), is a robust one, and is characterised by possessing
a nearly continuous spiral of zooecia, which is, however, broken up into groups by distinct
intervals. The spiral is arranged in such a way as to leave a considerable part of the thick
stem exposed to view. The stem is divided by stout diaphragms, perforated at their centre,
into internodes, each of which corresponds with a single group of zooecia. In rare instances a
diaphragm may fail to develop. The specimen 198. A. appears at first sight to have unusually
long internodes, since an interval of as much as 65 mm. may occur between two successive
1) Heller, C, 1867. "Bry. Adriat. Meeres", Verh. k. k. zool. -bot. Ges., XVII, p. 127, PI. V, figs 1, 2.
65
SIBOGA-EXPEDITIE XXVlIIa.
66
branches. But the occurrence of branch-scars and of diaphragms shows that the internodes are
really of the typical length ; and that branches have either failed to develop or have been lost.
When the stem has reached a certain age, a joint is formed in the region of the diaphragm,
and a similar joint occurs at the base of the branch.
The basal view (fig. 4) of the zooecia is very characteristic. The zooecia of the two
series are usually alternate, with interdigitating inner walls; but are sometimes opposite, the two
zooecia being then separated by a wall nearly at right angles to the proximal and distal walls.
At about the middle of the basal surface occurs a single communication-pore, by which the
funicular tissue of the zooecium is continuous with that of the stem. On the outer side of the
pore, the basal surface gives origin to a circular group of retractor muscles of the polypide.
In the retracted condition, in which the specimens are preserved, the muscle-fibres appear very
thick and short; being contracted to such an extent that the lophophore lies close to the basal
wall. In parts near the growing tips many of the young polypide-buds are so placed that their
lophophore is visible from its distal side (fig. 5); and the young tentacles are clearly seen to
be 8 in number. In this stage of development the polypide-bud lies close to the basal and
outer side of the zooecium, which has already acquired nearly its full length. The lophophore
lies in the same position as that described above for the fully developed, adult polypide, in
its state of complete retraction; and its plane is parallel to the surface of the parent-stem and
almost in contact with that surface. The caecum of the young stomach lies close to the basal
wall, projecting towards the line which separates the two series of zooecia. At a slightly latei-
stage the tentacles are seen to be elongating in the direction of the principal axis of the
zooecium; the young polypide still lying entirely in the basal part of the zooecium. A mass of
cells, the commencement of the vestibule, is differentiated at the distal end of the tentacle-
sheath ; and at first lies well within the proximal half of the zooecium, in a position far removed
from the future orifice, with which it soon becomes connected by a fine thread.
The zooecia in which the polypides are fully retracted are completely connate, as has
been described above. When the kamptoderm is everted a free cylindrical portion is formed,
as in the right hand zooecium of fig. 2. At the distal end of this is seen a comparatively
small collar. When degeneration of the polypide takes place, as in the next six zooecia of the
same figure, the everted kamptoderm, which has lost its vestibule and collar, forms a low
mammilla, marked by numerous circular cuticular striae. The form of the orifice, in zooecia
with a healthy polypide, is seen in the left part of the same figure, and is quadrangular, the
vestibule being about one third the length of the zooecium. The details of the alimentary canal
are not readily made out; but a distinct gizzard is present.
The zooecia, which are somewhat variable in length, are about 600 — 700 u. long. Their
basal wall, as shown in fig. 4, is from 200 to 300 u. in breadth, and from 80 to looy. in length.
In the specimens from Stat. 164(126. A., 507. B.) I have found a condition identical with
that described by Busk l) in the specimens which he referred to A. semispiralis Kirchenpauer.
An internode, distinguished as such by a diaphragm at each end, is shown in Busk's figs 3^, 36
1) Busk, G., 1886, Challengei Report, Pt L, p. 36, PI. VIII, figs 3, 3 a, 3/'.
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67
as hearing two or three groups of zooecia, instead of a continuous spiral. But the groups in
question may readily be regarded as parts of a single spiral. That this view is correct is proved
hy 126. A., in which, in one and the same branch, some of the internodes bear a single spiral,
of the form typical in the present species, while in other internodes the spiral is more or less
broken up, in various degrees. Thus in some cases the groups are separated from one another
by intervals of bare stem, while in others successive groups touch at their bases, being manifestly
parts of one spiral in which, here and there, two zooecia diverge from their bases, thus dividing
the spiral into discrete groups. In some of the branches of the last order, in which the limit
of growth has been reached, all the internodes bear these short discrete groups of zooecia.
The correctness of this conclusion has been confirmed by an examination of Busk's type-
*
material in the British Museum (87. 12.9.935,936), from the 'Challenger' Station 188, S. of
New Guinea. The spirit-specimen (936) has the same general appearance as that of the 'Siboga'
material. A large proportion of the internodes, both in older and in younger parts of the colony,
have the spiral broken up into 2 or 3 groups ; but the reference of the specimen to the present
species is indicated by the fact that in other parts the spiral is continuous and quite typical.
The condition figured by Busk as A. scmispiralis may thus probably be regarded as one which
is sometimes assumed by A. convoluta, under circumstances unfavourable to growth in full vigour.
Waters l) has described a similar specimen from Torres Straits. In the absence of figures I am
unable to express any opinion on the question whether " Serialaria [Vesicularia'ï) scmispiralis"
Kirchenp. ~) belonged to the same species or not.
Amathia convoluta Lamouroux and Serialaria convoluta Lamarck 3) were described in
the same year (18 16); neither author giving a reference to the other. Lamouroux stated,
however (p. vi) that in 1 8 1 2 he had named a part of Lamarck's collections for him ; and
Tenison-Woods, in the Memoir (p. 96) cited above, has used this as one of the reasons for
preferring Lamouroux' names to those of Lamarck, in cases where the same names were used.
MacGillivrav describes the spiral, in Victorian specimens, as being continuous but
"interrupted at the division of the branches". The figures given by this author seem to show
that the 'Siboga' specimens belong to the same species, although the groups of their successive
internodes are apparently always separated by a slight interval. It is clear that Busk, in the
'Challenger' Report, had wrongly ascribed the specimens recorded as A. spiralis to that species.
Mr Kirkpatrick *), who has for many years been in charge of the Polyzoa in the British Museum,
has pointed out that A. spiralis Busk, is probably A. convoluta Lamx; and MacGillivrav")
has come to the same conclusion.
Pergexs 6) has called attention to a series of unpublished Plates, by Desmarest and
Lesueur, of which the originals are said to belong to the Museum at Havre. The MS. describing
these Plates was deposited by the authors, in 1829. in the Library of the Muséum de Paris,
1 Waters, A. W., 1887; — see ref. in synonymy.
2) Kirchenpauer, G. H., 1869, "Museum Godeffroy. Cat. IV", Hamburg, p. xxxiv (Samoa).
3) This appears to be A. spiralis Lamx.
4) Kirkpatrick, R., 1888, "Pol. Port Phillip", Ann. Mag. Xat. Hist., (6) II, p. 19.
5) MacGillivrav. P. H., 1895, t. cit.. p. 133.
6) Pergexs, E. : — see synonymy. under A. crispa.
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68
where it is still preserved. From a copy of these unpublished Plates in my own possession I
am able to state that PI. XIII, fig. 4 of Desmarest and Lesueur gives what appear to be
very characteristic figures of the species under consideration. According to Pergens, the MS.
name of this species is Sirinx circumplicata \ and it is identified by him with A. crispa Lamarck.
It is unfortunate that the original names and descriptions were never published by Desmarest
and Lesueur ; but it seems obvious that most of the names published by Pergens in 1887 have
no validity, as they refer to figures which are not technically in existence, and for the most
part (as in the case under consideration) are mere citations of names, not accompanied by any
kind of description. It may be worth while to emphasize the view that even in cases where
Pergens has given valid reasons for retaining these MS. names, they cannot be ascribed a date
earlier than 1887, when they were first published; whereas Pergens appears to have assumed
that, after his introduction o'f them, they could be ascribed to the year (1829) when the MS.
was deposited in the Paris Museum.
2. Amathia disfans Busk. (PI. V, fig. 6, 7).
Atnathia distans Busk, 1886, Challenger Rep., Pt L, p. 33, PI. VII, figs 1 — 1 c.
? Amathia distans MacGillivray, 1895, "Austr. Species Amathia", Proc. R. Soc. Vict. (N. S.),
VII, p. 134, PI. C, figs 3, ia.
? Amathia distans Thornely, 1905, HERDMAN's "Rep. Pearl Oyster Fisheries G. Manaar*', Publ.
Roy. Soc, Suppl. Rep. XXVI, "Pol.", p. 128 (Gulf of Manaar; no description).
? Amathia distans Thornely, 1907, "Rep. Mar. Pol. Ind. Mus.", Ree. Ind. Mus., I, p. 196 (Coast
of Cheduba, Burma; Andamans; no description).
? Amathia acervata Lamouroux, 1824, "Encycl. Méthod.", "Zooph.", p. 45 (Japan).
? Amathia tor/nosa Waters, 1910 (nee Tenison-Woods), "Rep. Mar. Biol. Sud. Red Sea",
uBry. II", J. Linn. Soc., Zool., XXXI, p. 243, PI. XXIV, fig. 5.
a 1 -y TT \
' . ' \ Stat. 7. Near Reef of Batjulmati, Java, ie. Metres and more; coral and stones.
436. A. ) ' j j j
' "j Stat. 50. Bay of Badjo, W. coast of Flores, o — 40 Metres; mud, sand and shells.
284. A. Stat. 58. Anchorage off Seba, Savu, O — 27 Metres; sand. (On a stone).
371. C. Stat. 273. Anchorage off Pulu Jedan, Aru Islands, 13 Metres; sand and shells.
Zoarium small, repent, attached here and there to the substratum by rootlet-like tubes.
Internodes about 100 p. in diameter, with a thick cuticle. Zooecia biserial, in spiral groups
which occupy the distal part of the internode and usually form one complete turn or less. The
proximal part of the internode carries no zooecia, and about half the internode is thus bare.
The branching appears dichotomous, owing to the fact that the prolongation of the stem and
the lateral branch form nearly equal angles with the part of the stem preceding the bifurcation.
The ramification is not regularly alternate, a branch being sometimes developed on the same
side of the stem as its predecessor, but often growing out from a point which is 900 removed
from that which gives rise to the preceding and succeeding branches. Consecutive branches
thus commonly lie in two planes at right angles to one another, instead of lying, on opposite
sides of the stem, in the same plane, as in A. convohtta. A transverse diaphragm, sometimes
oblique, occurs at the distal end of the internode, in the region where the group of zooecia ends;
68
69
and the diaphragm at the base of the branch lies just on its proximal side. There are no
definite joints in the region of the diaphragms, although the stem may be somewhat constricted,
a short distance on the distal side of a diaphragm, both of the main stem and at the base of
the branch. Zooecia not completely connate, the distal two-fifths or more being free, and sub-
conical. Orifice terminal, obscurely quadrangular. Basal wall of the zooecium not conspicuously
broader than long, the zooecia thus not having the compressed form characteristic of those of
A. convoluia. Polypides with 8 tentacles and a gizzard.
The present species differs from A. convoluta in its smaller size, its repent habit, the
smaller diameter of its sterns and the shorter spirals formed by the groups of zooecia, and in
the fact that its zooecia are not completely connate. It is attached to its substratum, which is
sometimes a Sponge, by rootlets, which may be formed as a modification of an ordinary branch.
The branch thus modified usually assumes an irregular, sinuous outline, and gives off a new
rootlet here and there, a few diaphragms being developed.
The zooecia (with retracted polypides) reach a length of about 450 — 500 u.. The vestibule
is nearly half the length of the zooecium and the collar is small. In some of the old zooecia
the free terminal part is marked by numerous circular striae; a condition which probably
indicates the approaching degeneration of the polypide.
The discrimination of the species of Amathia is well known to be difficult; and there
is much uncertainty with regard to the synonymy. The present specimens appear to be
indistinguishable from the type-specimen (Brit. Mus., 87. 12. 9. 926) of A. distans, described by
Busk l) from Bahia, Brazil. With that species it agrees in its straggling growth, in the restriction
of the spiral groups of zooecia to the distal half of the internode, in the fact that the zooecia
are only partially connate, and in the measurements. The internodes are described by Busk as
being 150 u. in diameter, and the zooecia are 400 — 500 u. in length. A South Australian form
has been described by MacGillivray -), under the same name. The species recorded by Waters 3)
from the Sudanese Red Sea, under the name of A. fortuosa, is similar to the 'Siboga' specimens
in the form of its zooecia, but its internodes seem to have a thin cuticle. I venture to doubt
the correctness of Mr Waters' determination, since the zooecia are shown as very short and
somewhat discrete; whereas in the original description of Tenison-Woods, the "great length of
the pairs of cell" *) is emphasized, and they are represented as completely connate; the spiral
being, moreover, much elongated and occupying most of the length of the internode. A. distans,
mentioned by Waters in the same paper, is figured (PI. XXIV, fig. 7) as having connate zooecia.
The 'Siboga' specimens show considerable resemblance to A. pruvoti, described by
Calvet 5) from Banyuls-sur-Mer; but they differ from that form in the fact that the zooecia
are less completely connate.
1) Busk, G., 1886, loc. cit.
2) MacGiu.ivray, P. H., 1895, loc. cit.
3) Waters, A. W., 1910, loc. cit.
4) Tenison-Woods, J. E., 1880, "On the Genus Amathia", Trans. Proc. R. Soc. Vict., XVI, p. 89. 2 figs-on unnumbeied Plate.
In Tenison-Woods' fig. 6, the zooecia appear rather short. It is possible that the words "the great length of the pairs of cells in each
internode" were intended to refer to the length of the spiral group : — which would be more in accordance with the figure.
5) Calvet, L., 1911, "Deux Esp. Nouv. Bry. Mediterranée", Arch. Zool. Exp. (5) VIII, Notes et Revue, p. I.IX, text-fig. on p. lx.
69
yo
A. brasiliensis Busk (1886, p. 34, PI. VII, figs 2 — 2e) resembles A. distans in many
of its characters, but is a more robust species, its sterns being about twice as broad as those
of A. distans. The respective diameters are given by Busk as '3 mm. and -15 mm.
It is not impossible that the present species is identical with A. accrvata, described by
Lamouroux (1824) from Japan.
Bowerbankia Farre.
Bowerbankia Farre, 1837, Phil. Trans., p. 391.
Bowerbankia Hincks, 1880, "Hist. Brit. Mar. Pol.", p. 51S.
Bowerbankia Waters, 1910, "Rep. Mar. Biol. Sud. Red Sea", "Bry. II", J. Linn. Soc. Zool.,
XXXI, p. 240.
Zooecia arising irregularly from an erect or creeping axis, commonly in definite groups.
Tentacles 8 — 10. Gizzard present.
Mr Waters1) regards Bowerbankia as a synonym of Zooèotryou Ehrenberg, 1831. In
this conclusion I am not at present prepared to follow him ; as it seems to me that the zoarial
characters are sufficiently distinct to warrant the separation of the two genera. I am, however,
in agreement with him in thinking that Bowerbankia biserialis Hincks 2) is a synonym of Z.
pellucidum ; that Vesicularia bilateralis MacGillivray 3) and an unnamed form described by
Macdonald 4) belong to the same genus, if not to the same species ; and that Dedalaca
manritiana Quoy and Gaimard 5) is probably also referable to Zoobotryon, as was suggested
by de Blainville °).
It may be noted that "Zoobotryon sp." has been recorded by Whitelegge 7) from the
neighbourhood of Sydney ; and that Z. pellucidum has been described by Miss Philipps s) from
the Isle of Pines, New Caledonia. From the Atlantic region the same species has been recorded
from Brazil (F. Muller, 1860), Florida (Osburn, 1914) and Bermuda (Smallwood, 19 10). In
spite of its wide distribution I have not, however, found it among the 'Siboga' dredgings.
1. Boiverbankia imbricata Adams 9) (? sp.). (PI. VII, figs 15, 16).
Sertularia imbricata Adams, 1800, "Descr. Mar. An. Wales", Trans. Linn. Soc., V, p. 11,
PI. II, figs 5— 11.
Bowerbankia imbricata Waters, 1910, "Rep. Mar. Biol. Sud. Red Sea", "Bry. II", J. Linn.
Soc. Zool., XXXI, p. 248, PI. XXV, figs 6—10.
1) Waters, A. W., 1910, t. cit., pp. 240, 244.
2) Hincks, T., 1887, Tol. Adriat.", Ann. Mag. Nat. Hist. (5) XIX, p. 309, PI. IX, figs 6, 6 a.
3) MacGillivray, P. H., 1889, "S. Austr. Pol.", Trans. Proc. R. Soc. S. Austr., XII, p. 30, PI. II, fig. 4.
4) Macdonald, J. D., 1857, "Brief Descr. Cten. Pol.", Proc. Roy. Soc, VIII, p. 383 (reprinted, under the same title, in Ann.
Mag. Xat. Hist. (2) XIX, 1857, p. 390).
5) QüOY and Gaimard, "Voy. Astrolabe", "Zool.", IV, 1834, pp. 299 — 301; III, 1835, p. 952 (expl. of Plate); Atlas, "Zooph.",
PI. XXVI, figs 1, 2, 21. For dates of publication see Siiert.orn and Woodward, Ann. Mag. Nat. Hist. (7) VIII, p. 333.
6) Blainville, H.M.D. de, 1834, "Man. Actinol.", pp. 679, 680; see also pp. 493, 494, PI. LXXXI, figs 6, 6a. De Blainville's
actual words are "Il faut sans doute rapprocher de ce genre (Dctfalaea) celui que M. Ehrenberg nomme Zooiotrion".
7) Whitelegge, T., 1889 (1890), p. 293.
8) Philipps, E. G., i899,pp. 441, 450.
9) The description and figures of this author are hardly recognisable ; but in view of the desirability for stability in nomenclature,
it seerns to me best to assume that Adams' species was the one generally known as Bowerbankia imbricata.
70
7i
Bowerbankia densa Farre, 1837, t. cit., p. 391, Pis XX, XXI.
Bowerbankia imbricata (incl. B. densa) Hincks, 1880, t. cit., p. 519, PI. LXXIII, figs 1, 2.
? Bozverbankia imbricata, form densa Hincks, 1884, "Pol. Q. Charlotte Is", Ann. Mag. Nat.
Hist. (5) XIII, p. 207.
'? Stat. 50. Bav of Badjo, W. coast of Flores, o — 40 Metres; mud, sand and shclls.
325- J- '
382. B. Stat. 64. Kambaragi Bay, Tanah Djampeah, O — 32 Metres; coral, coral-sand.
338. B. Stat. 71. Makassar, o — 32 Metres; mud, sand with mud, coral.
' ' > Stat. 164. i°42.s S., I30°47'.5 E., 32 Metres; sand, small stones and shells.
374. K. (No locality recorded). On Peropliora.
Japan, Mus. Zool., Cambridge, Reg. June 23, 1902 (A. OwsTON Coll., 27. D., off Tokyo,
150 fathoms). (On Bugula johnstoniaè).
Characters of the material here recorded: —
Zooecia arising in small groups from a creeping stolon ; sometimes less definitely grouped
or even arising singly. The stolon is occasionally septate, and gives off a few branches here
and there. Zooecia elongate, subcylindrical, straight or somewhat curved, usually paired, their
basal ends much wider than the stolon, sometimes produced into processes, and projecting on
either side of it. Orifice square. Vestibule rather long, but variable in length ; containing a
moderately developed collar. Tentacles not numerous, probably not more than 10. Pharynx
large, oesophagus short, bent during retraction, passing into a short conical proventriculus,
which is succeeded by a well marked gizzard. Stomach and caecum large; intestine and rectum
narrow. Parietal muscles extending along the whole length of the zooecium, the fibres usually
arranged singly.
I think I am right in referring all the specimens above recorded to a single species;
though some of them are obscured by the remains of Sponges in which they were more or
less embedded. As in many other species of Ctenostomata, the zooecia vary much in length ;
but, as in other cases, this appears to depend largely on changes accompanying the histolysis
of the polypide. The best specimen, part of which is shown in figs 15, 16, is 374. K., the
locality of which, by some oversight, was not recorded. It is growing on the oral end of a
Perophora, to which the stolon is for the most part adnate. The complete zooecium shown in
fig. 16 is 980 y. long and 190 u. in diameter. The longest zooecium in fig. 15 is 1200 u. in
length ; but its vestibule has lost its connexion with the rest of the polypide, preparatory to
histolysis, and the collar is completely protruded. The cuticle is very thin and transparent. The
vestibule is provided with four groups of parieto-vaginal muscles, close to its proximal end ; and
there are four groups of similar muscles, close to the first set, but inserted into the junction
of the vestibule and tentacle-sheath.
Several of the specimens show occasional diaphragms, traversing the stolon; but I am
not able to state the typical arrangement of these structures.
The specimen from Japan appears to belong to the same species; but the preparation
is not a very successful one. Hincks l) has alluded to "a Vesicularian, apparently allied to
1) Hincks, T., 1880, t. cit., p. 519, note.
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72
B. pustit/osa, which had been brought by Dr v. Martens from Yokohama", and had been
recorded by Kirchenpauer *).
I think there is a high probability that the 'Siboga' specimens belong to the species
which has been recorded by Waters, from the Red Sea, as B. imbricata, although I am not
certain that his determination is correct. The specimens under consideration resemble those
figured by Waters, but I cannot believe that the alimentary canal is correctly represented in
fig. 8 of that author. It may also be identical with the B. caudata described by Annandale 2)
from Indian localities. But Waters does not admit that B. caudata is a good species, and
points out that the tail-like process at the basal end of the zooecium is not distinctive, as it
also occurs in some individuals of undoubted B. imbricata. The specimen (374. K.) figured
in the present Report has no basal processes, which are, however, present in other 'Siboga'
specimens (325. J., 382. B., 130 M.) ; either as a pointed prolongation of the kind figured by
Annandale, or more commonly as several irregular processes,. like those shown by Waters in
his PI. XXV, fig. 10 ; and regarded by him as rooting structures by means of which the
zooecium is attached to its substratum. B. imbricata, according to the same author (p. 249)
has the largest gizzard of any species with which he is acquainted, its transverse diameter being
about 100 a. This is in fair agreement with my own measurement of the gizzard of the specimen
shown in fig. 16; the transverse diameter of which I find to be 90 u.. If the number of tentacles
in the 'Siboga' specimens is 10, as I think is probably the case, this is a further agreement with
B. imbricata-, some of the other species of the genus being known to have only 8 tentacles.
Miss Robertson 3) has recorded B. imbricata from the American side of the Pacific;
but she has given no description of the specimens found there.
Group D. Stolonifera Ehlers.
Stolonifera (pars), Ehlers, 1876, "■Hy pophor ella expansa' , Abh. K. Ges. Göttingen, XXI, p. 126.
Stolonifera Waters, 1910, "Rep. Mar. Biol. Sud. Red Sea", "Bry. II", J. Linn. Soc. Zool.,
XXXI, p. 241.
Waters, 1. cit., has proposed a new grouping of the genera formerly placed together as
Stolonifera; and the distinctive characters of the Stolonifera, as thus restricted by him, have
been stated above, on p. 43. The axis, in this group, is delicate, and thus contrasts with
the more massive axis of the Vesicularina. It expands slightly here and there, a diaphragm
being formed immediately on the distal side of the expansion, which gives rise either to new
stolon-branches or directly to zooecia, which are usually arranged in pairs. A gizzard is
usually absent.
1) The reference to Kirchentauer's Memoir, which is not given by HlNCKS, is "Zoolog. Ergebn. Nordseefahrt", VI. "Bryozoa",
II Jahresb. Komm. Cnt. deutsch. Meere Kiel, 1874, p. 192.
2) ANNANDALE, N., 191 1, "Fauna of Brit. India", "Freshwater Sponges, Ilydroids and Polyzoa", p. 189. See also the reference
to Miss T110RNEI.Y, there given.
3) Robertson, A., 1900, "Papers Harriman Alaska Exp.", Proc. Washington Acad. Sci., II, p. 331. In "Non-incrnst. Cheil.
Bry. W. Coast N. Amer." (Univ. Cal. Publ. Zool., II, N° 5, 1905, p. 239) she figures "/?. imbrkata" with 8 tentacles, a fact which throws
poubt on the correctness of the other determination.
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73
Fam. Valkeriidae Hincks.
Valkeriidae Hincks, 1S77, "On British Polyzoa", I, Ann. Mag. Nat. Hist. (4) XX, p. 532.
Valkeriidae Hincks, 1S80, "Hist. Brit. Mar. Pol.", p. 551.
The Family Valkeriidae was defined by Hincks in 1880, as the first of his group
Campylonemida, which he supposed to be characterised by having two of the tentacles everted
when protruded. The validity of this character has been criticized by other writers1); and the
existence of two everted tentacles cannot at any rate be ascertained in most preserved specimens,
in which the tentacles are usually retracted. Hincks defines the Family as having "Zooecia
contracted below, deciduous, destitute of a membranous area". The existence of 8 tentacles and
the absence of a gizzard were given as characteristic of the Campylonemida in general.
Valkeria Fleming.
Walker ia Fleming, J., 1823, Wern. Mem., IV, Pt II, p. 490.
Valkeria (pars), Fleming, 1828, "Hist. Brit. An.", p. 550.
Valkeria Farre, 1837, Phil. Trans., p. 402.
Valkeria Hincks, 1880, t. cit., p. 551.
Valkeria Waters, 1910, t. cit., p. 241.
Zooecia ovate or cylindrical, narrow at their proximal end and deciduous, arising from
the sides or from the distal end of an internode. Tentacles 8. Gizzard wanting. Orifice more
or less quadrangular.
The present genus was named after Dr Walker, and appears as Walkeria in Fleming's
original account (1823). In his later work (1828) Fleming uses the form Valkeria without even
mentioning the fact that it had at first been spelt differently. Valkeria has been universally
adopted by later writers, and it appears to me most undesirable to revert to the original spelling.
1. Valkeria atlantica Busk. (PI. VI, figs 5 — 12).
Farrella atlantica Busk, 1886, Challenger Rep., L, p. 7,/t PI. VII, figs 3 — $g.
? Farrella atlantica Thornely, 1905, Herdman's "Rep. Pearl Oyster Fisheries G. Manaar",
Suppl. Rep. XXVI, publ. Roy. Soc, p. 128 (110 description).
? Farrella atlantica Thornely, 1907, "Rep. Mar. Pol. Ind. Mus.", Ree. Ind. Mus. I, p. 196
(Burma; no description).
? Farella atlantica Thornely, 1912, "Mar. Pol. Ind. Ocean", Trans. Linn. Soc. 2 Ser., Zool.,
XV, p. 157 (no description).
' ' > Stat. 7. Near reef of Batjulmati, Java (from reef). On 435. A., Farcimia; etc).
380. G. Stat. 47. Bay of Bima, 55 Metres; mud with patches of fine coral-sand. (Growing in
profusion on Hydroids.)
20. E. )
23. A. Stat. 50. Bay of Badjo, W. coast of Flores, o — 40 Metres ; mud, sand and shells.
325. M.)
1) As by Loppens, 1908, "Rry. d'eau douce", Ann. Biol. lacustre, III, p. 149.
73
SIBOGA-EXPEDITIE XXVIII Cl.
74
'' f Stat 71. Makassar, O — 32 Metres; mud, sand with mud, coral. (On Scrupocellaria, etc).
338. C. )
92. F. Stat. 117. Kuandang Bay, 80 Metres; sand and coral. (On Peropliora).
393. C. Stat. 133. Anchorage off Lirung, Salibabu Island, o — 36 Metres; mud and hard sand.
(On a Perforate Coral).
108.O. Stat. 144. Anchorage N. of Salomakiëe (Damar) Island, 45 Metres; coral-bottom and
Lithothamnion.
396. R. Stat. 162. Between Loslos and Broken Islands, W. coast of Salavvatti, 18 Metres;
coarse and fine sand with clay and shells.
501. H. Stat. 163. Anchorage near Seget, W. entrance Selee (Galewo) Strait, 29 Metres; sand
and stone, mixed with mud. (On Scrupocellaria).
? Stat. 164. i°42'.5 S., i30°47'.5 E., 32 Metres; sand, small stones and shells.
139. F.)
445. B. ] ^ " | Ambon Anchorage, 36 — 54 Metres; mud, sand and coral. (On Caberea).
348. B. Stat. 213. Saleyer, O — 36 Metres; coral-reefs, mud and mud with sand.
296. C. Stat. 273. Anchorage off Pulu Jedan, Aru Islands, 13 Metres; sand and shells.
415. D. Stat. 318. 6°36'.5S., U4°55'.5E., 88 Metres; fine yellowish grey mud. (On 415. C,
Carbasca cribriformis).
Zoarium delicate, typically repent, but sometimes becoming detached from its substratum.
Stolon slender, branching in a cruciform marmer, consisting of a succession of internodes, of
very varying length, separated from one another by diaphragms. Branches given off at the distal
ends of the internodes, immediately before a diaphragm, and separated from the parent-internode .
by a lateral diaphragm. The lateral branches may consist of a simple series of internodes or
may give rise to new branches. Zooecia arising from a vestigial internode, which may either
replace a lateral branch or may occur immediately on the distal side of the base of a branch.
Proximal end of the zooecium slender, stalk-like, usually short but sometimes greatly elongated.
The rest of the zooecium is elongated, subcylindrical and relatively slender. Orifïce terminal,
subquadrangular ; vestibule rather short ; collar small and delicate. The histolysis of the polypide
is foliowed by a great shortening of the zooecium, the body-wall of which becomes marked by
numerous circular striae. Polypides with 8 tentacles, which may be bent at their tips during
retraction. Gizzard wanting.
I can find no differences between the specimens here recorded and the species described
by Busk as Farrella atlantica, from Bahia, Brazil. The 'Siboga' material has been compared
with the type-specimens of Busk's species preserved in the British Museum (Slide 99. 7. 1. 432CN
on Amathia brasiliensis; and unmounted material, on the same species, 87. 12.9.928; and on
A. distans, 87. 12.9.926). The 'Challenger' specimens show the same differences in the form
of the zooecia, in different states, that are here described. If the determination is correct it
may be concluded that the present species has a wide distribution ; and that it will probably be
discovered in many intermediate localities. It is probably the species which has been recorded,
without description, by Miss Thornely, from several localities in the Indian Ocean.
Waters1) has recorded F. atlantica from Zanzibar and British East Africa; but the
muscles described at the proximal end of the zooecium appear to indicate that this species
should be referred to Mimosclla.
1) Waters, A. W., 1914, p. S52, PI. IV, flg. 9.
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75
It is very doubtful whether Bcsk's species was correctly placed in Farrella l), which is
characterised, according to the diagnosis given by Hincks 2), by having a bilabiate orifice.
F. repens is said by the same author to have 10 — 12 tentacles.
From V. uva the present species seems to be separated by the mode of origin of the
zooecia. The arrangement is both characteristic and constant, a single zooecium or a pair of
zooecia being developed at the distal end of an internode, and ahvays from a minute rounded
or slightly pointed structure (figs 5 — 7) which appears to be a vestigial internode. The zooecia
may be unilateral or ma}- occur on both sides of the internode, sometimes with the addition
of a lateral branch. In V. uva the distal end of the internode may give rise to several zooecia
on each side.
In most of the specimens a considerable proportion of the zooecia have assumed the
contracted form shown in figs 5, 6. This change, which has not escaped the notice of previous
writers (e.g. Waters3)), and has already been alluded to in the description of several of the other
Ctenostomes recorded in the present Report, is associated with the degeneration of the polypide.
The shortened zooecia are usually more or less pear-shaped, but may assume irregular shapes
(fig. 6). When fully formed they have no obvious contents except a brown body, or sometimes
two of these structures. Fig. 5 shows that a new polypide may be developed in a zooecium
which has undergone this change.
The zooecium contains no definite muscles by which the contraction could be effected;
and the occurrence is perhaps due to a loss of turgidity, by the absorption of some of the
water contained in the body-cavity. The process is probably aided by the contraction of the
cuticle, which may be supposed to be elastic and somewhat stretched in the ordinary condition.
The circular markirig of the body-wall which is a characteristic feature of zooecia in this
condition is satisfactorily accounted for on this hypothesis.
The zooecia in their active condition (fig. 7) have a form very different from that of
those just described. They are elongated, straight or curved, and are more or less cylindrical,
though frequently dikited in their proximal half. They have a very transparent body-wall, which
is well stretched and has none of the circular markings of the contracted zooecia. At their
proximal end they taper off into a stalk-like portion, which is usually short but in the specimen
393. C. (figs 9 — 12) is much elongated.
The vestibule is relatively short, and the collar is very delicate and small. The tentacles
are 8 in number, and are commonly bent at their tips during retraction ; in which condition
the oesophagus is folded on the pharynx. There is no gizzard; in which respect the present
species agrees with the usual definition of the genus.
Some of the zooecia (e. g. in 337. B.), of the ordinary, uncontracted form are fertile.
The vestibule persists, but the rest of the polypide has atrophied, with the exception of a sac
which encloses the embryo and is no doubt the remains of a tentacle-sheath. The embryo is
1) Farrella Ehrenberg, 1839, Üb. d. Bildung d. Kreidefelsen", Phys. Abhandl. k. Ak. Wiss. Berlin, a. d. Jahre 1838, Table II
(= new name for Lagcnclla Karre, 1837, preoccupied by Lagc/itlla Ehrb., 1832).
2) Hincks, 1880, t. cit., p. 52S.
3) Waters, A. W., 1910, p. 239.
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;6
solitary and is elongated in the same direction as the zooecium, reaching about half its length
and about three quarters of its width. An advanced embryo, to which this description refers,
measures 205 by 70 a.
The zooecia vvith functional polypides are 600 — 650 p. in length and 70 — 85 p. in greatest
breadth (fig. 7); but the elongated zooecium shown in fig. 10 is 1140 ij. long. In other cases
the length may not exceed 450 ij.. The contracted zooecia shown in fig. 5 are 220 — 250 ft in
length and 70 — 90 u. in breadth. It may be noticed that the stalk-like proximal portion of
these individuals remains uncontracted. The stolon is about 1 9 u wide.
This very beautiful species ma)- probably be regarded as the commonest Ctenostome in
the area investigated, since I have found it from fourteen of the Stations, ranging from Java
in the West to the Aru Islands in the East and Lirung in the North; the triangle defined
by these three points including nearly all the Stations at which Polyzoa were found. It is so
inconspicuous that it is not readily seen except by microscopic examination ; and it has thus
always been found accidentally, on some other species which has been large enough to have
been recognised as belonging to the collection of Polyzoa. It is probable that an examination
of the specimens of Hydroids and other objects obtained by the Expedition would result in its
discovery from many Stations where I have not found it.
2. Valkeria tuöerosa Heller. (PI. VI, figs 13 — 20).
Valkeria tuöerosa Heller, 1867, "Bry. Adriat.", Verh. zool. -bot. Ges. Wien, XVII, p. 129,
PI. VI, figs 3, 3«.
Valkeria tuberosa Carus, 1889, "Prodr. Faun. Med.", II, p. 52.
Valkeria tuberosa Vine, 1891, "Brit. Pal. Cten. Pol.", Proc. Yorksh. ■Geol. Polyt. Soc, XII,
p. 91, PI. IV, fig. 2 (recent, Naples; figured for comparison with fossil species).
Valkeria uva (pars), Waters, 1910, "Rep. Mar. Biol. Sud. Red Sea", "Bry. II", J. Linn. Soc.
Zool., XXXI, p. 250, PI. XXIV, fig. 13 (Naples).
332. B. Stat. 64. Kambaragi Bay, Tanah Djampeah, o — 32 Metres ; coral, coral-sand. (On
332. A., Micropora ratoniensis Waters).
57. R. Stat. 80. Borneo Bank, 40 — 50 Metres; fine coral-sand.
488.I. Stat. 133. Anchorage off Lirung, Salibabu Island, o — 36 Metres; mud and hard sand.
'I Stat. 164. i°42'.5 S., I30°47'.5 E., 32 Metres; sand, small stones and shells. (On other
" t Polyzoa, Hydroids, Alcyonaria, shell, etc).
360. C. Stat. 240. Banda Anchorage, 9 — 45 Metres; black sand; coral, Lithothamnion-bank.
255. D. Stat. 311. Sapeh Bay, E. coast of Sumbawa, o — 36 Metres; mud and sand.
Zoarium repent, its narrow stolon attached to Polyzoa, Hydroids, Alcyonaria, shells and
other objects. The stolon dilates at intervals into a slight swelling, cut off distally by a diaphragm
and producing, on each side, a branch composed of short internodes, each of which gives off
paired lateral branches like those of the main stem. Zooecia and new stolons are produced
by the branches thus formed. Owing to this mode of branching and to the shortness of the
internodes, the zooecia are crowded into a group which is circular in outline when seen from
above. Zooecia subcylindrical, narrower at the base, of nearly uniform width, and relatively
broad and short. Orifice terminal, more or less quadrangular. Everted kamptoderm considerably
76
77
narrower than the zooecium and, like the vestibule, about half the length of the fully retracted
zooecium; bearing a well developed collar. Tentacles 8, bent distally during retraction. Oeso-
phagus long, folded on the pharynx when the polypide is retracted. Gizzard wanting.
This species was originally described by Heller from the Adriatic. I should hardly have
ventured to refer the 'Siboga' specimens to it if it had not been for the evidence of specimens
from Naples1) in the collections of the British Museum (88. ii. 9. 38) and the University Museum
of Zoology, Cambridge. These agree so closely with the Malay specimens, even in the character
of the branching, that I am unable to detect any differences of importance. The groups of
zooecia shown by Heller in PI. VI, fig. 3 have a close resemblance to those figured in the
present Report. The lateral internodes are represented by Heller as being almost spherical,
although the Xeapolitan specimens agree closely with those from the 'Siboga' collection in their
form. When Heller describes the zooecia as somewhat smaller at their distal end (his fig. 3«)
he is 110 doubt alluding to zooecia like those shown in my own figs 19, 20.
The stolon of the Malay specimens is narrow, measuring about 45 u. in diameter; and
the older parts have a thick, chitinous wall. The mode of branching is represented in fig. 15,
from which it will be seen that it is identical with that of a specimen from Naples which has
been figured by Waters 3). The parent-internode is marked off by a distal diaphragm, but there
is 110 structure of this kind at the proximal end of the dilated portion. It gives off a pair of
lateral branches, which in the specimen figured are each composed of two short internodes,
prolonged into a stolon distally. Each of these internodes gives off a pair of lateral branches
of similar constitution. The distal internode of each secondary branch commonly gives rise to
(a) a distal stolon, (ó) a pair of zooecia, (V) a pair of lateral stolons situated just proximally to
the zooecia. This arrangement is, however, not constant, and a median zooecium may be formed
distally in addition to a distal stolon, which comes off on the basal side of the zooecium. Other
zooecia are given off by the other internodes of the system.
In fully developed groups of zooecia (fig. 1 3) it is usually difficult to ascertain the details
of the branching, which may perhaps be more complex than in the specimen shown in fig. 15.
In these groups the zooecia vary in form according to their state of development. The young
zooecia are almost spherical (figs 14, 15); but the old ones, with functional polypides, have
the form which has been described above as typical (figs 13, 16).
In a considerable number of cases the zooecia have died with the kamptoderm everted,
as shown in fig. 17. This region is less than half the width of the rest of the zooecium, and
is terminated by the collar, which is conical in form and has strongly marked ribs or folds.
Zooecia in this condition usually show some indication of degeneration of the polypide, the first
evidence of which is seen in the degeneration of the vestibule. There seems reason to believe
that the collar is thrown off after this stage has been reached : — a process which I believe
to take place commonly in Ctenostomata. In a later stage, when the histolysis of the polypide
has advanced, the zooecia have the form represented in figs 19, 20 and also in Heller's fig. 3^.
The everted kamptoperm has become short and thick walled, and is usually marked by circular
1) Cf. also Vine, 1. c.
2) Waters, A. W., 1910, 1. c.
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78
cuticular striae. Many of these zooecia are practically empty of cellular contents except for one
or two brown bodies: but in sonie of them a young polypide-bud can be seen close to the
basal end of the zooecium. It may thus be inferred that in a certain proportion of cases the
polypide is reconstituted in zooecia which have undergone histolysis.
The structure of zooecia with functional polypides is shown in fig. 16. The quadrangular
orifice leads into a vestibule which is of great length, as it is about half the length of the
zooecium. The lophophore lies near the basal end of the zooecium during retraction, and the
tentacles are bent at their tips. Their number (8) could be ascertained with certainty from one
or two polypides which had died with their tentacles protruded. The pharynx is large and lies
in the prolongation of the line of the bundie of tentacles. The oesophagus commences close to
the proximal end of the zooecium and runs distally. It is of considerable length and it opens
into the stomach without forming a gizzard. The parietal muscles consist of a doublé series of
sharply defined bundies, each of two or three strong fibres.
The zooecia contain a number of minute, sharply defined spherical vesicles, varying in
diameter from 3.5 to 1 1 [j.. These structures appear to be of the same nature as the "excretory
vesicles" which I have described in certain Cheilostomata l) and Cyclostomata 3).
The following measurements, in y., will give an idea of the size of the zooecia and other
parts: — length of zooecium with retracted polypide, 700 — 850 ; diameter, 200 ; length of
vestibule or everted kamptoderm, 350 — 400; of collar, 210; diameter of stolon, 40 — 50.
Fam. Mimosellidae Hincks.
Mimosellidae Hincks, 1877, "On Brit. Pol.", II, Ann. Mag. Nat. Hist. (4) XX, p. 532.
Mimosellidae Hincks, 1880, "Hist. Brit. Mar. Pol.", p. 555.
This Family, which contains the single genus Mimosella, is distinguished by possessing
a pair of muscles by which the main part of the zooecium is moveable on its narrow stalkdike
proximal portion.
Mimosella 3) Hincks.
Mimosella Hincks, 185 1, "Notes Brit. Zooph.", Ann. Mag. Nat. Hist. (2) VIII, p. 359.
Mimosella Hincks, 1880, 1. cit.
Mimosella Waters, 1910, "Rep. Mar. Biol. Sud. Red Sea", "Bry. II", J. Linn. Soc. Zool.,
XXXI, p. 241.
Zoarium erect or repent. Zooecia deciduous, narrow proximally, in which region there
are paired muscles by means of which the zooecium can be moved on its stalk. Tentacles 8.
Gizzard wanting 4).
1) Harmer, S. F., 1891, "Excr. Processes Mar. Pol."', Q. J. Mier. Sci., XXXIII. p. 129.
2) Ibid., 1898, "Dev. Tubulipora\ Q. J. Mier. Sci., XLI, p. 113.
3) Mr E. O. ULRICH has suggested that l'inclla. a supposed Lower Silurian Ctenostomatous Polyzoon, is related to Vesicularia
and probably to Mimosella (1890, "New Lower Silurian Bry.", J. Cincinnati Soc. Nat. Hist. XII, p. 173). See also Clrich and Bassler,
1904, "Rev. Pal. Bry.", I, "Ctenostomata", Smithson. Misc. Coll. (Quarterly issue) XLV, pp. 256 et seq.
4) Vigelius, W. J., (1887, "Zur Morph. d. mar. Bry.", Zool. Anzeiger, X, p. 239) states that a gizzard is present in M. gracilis\
but Waters (t. cit., p. 246) denies the existence of this structure in Mimosella.
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79
i. Mimosella bigeminata Waters. (PI. VII, rïgs i — -7).
Mimosella bigeminata Waters, 1914, "Mar. Fauna Brit. E. Afr.", "Bry. Cycl. . . .", Proc. Zool.
Soc, p. 851, PI. III, figs 1 — 3 (Zanzibar Channel, 10 fathoms).
38. M. Stat. -j-j. Borneo Bank. 3°27'S., 117° 36 E., 59 Metres; fine grey coral-sand.
396.O. Stat. 162. Between Loslos and Broken Islands, W. coast of Salawatti, 18 Metres;
coarse and fine sand with clay and shells.
123, A. Stat. 164. i°42'.5 S., I30°47'.5E., 32 Metres; sand, small stones and shells.
359. C. Stat. 240. Banda Anchorage, 9 — 36 Metres; black sand, coral, Lithothamnion-bank.
374. D. Locality and depth not recorded.
Zoarium Stirparia-\\ke, consisting of much elongated, attenuated, unbranched sterns rising
from a basal series of rootlets or stolons. Erect sterns jointed, commencing with a few barren
segments, follovved by one or tvvo with a single pair of zooecia ; the remaining stem-segments
bearing two pairs of zooecia at their distal end. Zooecia long, subcylindrical, provided with a
proximal cone of muscles by which they are no doubt moved on the bracket-like support from
which they originate. Vestibule short, with a very delicate collar. Tentacles 8. Alimentary canal
not folded during retraction. A slight dilatation of the alimentary canal, at the junction of the
oesophagus with the stomach, may be the morphological equivalent of a gizzard; but a structure
actually of this nature can hardly be said to be present.
There can be little doubt of the corrrectness of the reference of this species to Mimosella,
a genus which has the peculiarity of having its zooecia moveable on their stalk-like proximal
ends. Hixcks has given a good description of these movements, from observations made on
living specimens of M. gracilis l) ; and he compared the movements of the zooecia with those
of the leaflets of Mi?//osa, the Sensitive Plant. He was unable to find any special muscles *),
which could be regarded as the agents of these movements. In the 'Siboga' specimens it is
easy to recognise these muscles in a group of cone-like form, situated at the extreme proximal
end of the zooecium. The muscles in question originate in the bracket-like projection which
supports the zooecium, and diverge to be inserted into the inner side of the body-wall of the
zooecium (fig. 3). In suitably orientated specimens it can be seen that the cone consists of a
pair of groups of fibres. Hincks' observation that the movements of the zooecia continue after
the loss of the polypide is accounted for by the fact that the muscular cone does not disappear
when histolysis of the polypide takes place. The movements may perhaps be affected by a
special arrangement of muscles, described below, in the stem-segments.
The general characters of the species may be illustrated by a description of the specimen
shown in hg. 5. The erect stem originates from an internode of the creeping stolon from which
rootlets diverge in various directions over the substratum. The stem is divided into segments
by diaphragms; and at each such point it is jointed by a slight annular thinning of the cuticle,
which is elsewhere sufficiently developed to give the stem-segments a rigid character. The first
stem-seo-ment is lono- (2160 u.) and bears no zooecia. It is foliowed bv three other barren
1) Hin-cks, T., 1851, "Notes Brit. Zooph.'', Arm. Mag. Nat. Hist. (2) VIII, p. 359; and 1880, "Hist. Brit. Mar. Pol." pp. 556, 558.
2) They have, however, been figured by Joliet (i888, "Études Pyresoma Rech. Bry. Roscoff et Menton'\ Paris, p. 112,
PI. V, fig. 5 wc) and by Waters (1914, t.cit., PI. III, fig. 1).
79
8o
segments, which diminish successively in length. The next segment bears a pair of brackets,
representing the bases of the distal zooecia of the more typical stem-segments ; but the zooecia
themselves, as in the next following segments, have fallen off. The next segment, and all its
successors, bear two pairs of brackets, the proximal pair being situated more laterally than the
distal pair. Between the proximal brackets the cuticle has an oval fenestra, closed by membrane,
into which are inserted the ends of a linear series of short muscle-fibres l), which originate in
the opposite wall of the stern-segment (figs 3, 4). It seems clear that the contraction of these
fibres must increase the pressure of the fluid in the cavity of the segment. This presumably
increases the turgidity of the segment, and probably makes it sufficiently rigid to withstand the
strain caused by the contraction of the muscles which move the zooecia. Occasional fenestrae
of more irregular form may be seen on other parts of the wall of the stem-segments.
Fifteen stem-segments have been omitted, and the upper part of the figure shows the
remainder of the stem. The deciduous zooecia have disappeared from all the segments omitted
in the figure, with the exception of one which bears a single zooecium, and from the two lower
segments of this upper part of the stem ; and some of the succeeding segments do not retain
the full number of zooecia. But the constant occurrence of two pairs of brackets may be taken
as a sure indication that each of these segments has borne its full number of zooecia at some
time during the growth of the stem. Xear the distal end, each segment has four zooecia.
The typical arrangement can be seen more clearly in figs 1, 2, showing the growing
ends of two sterns under a higher magnification. The zooecia originate near the tip of the stem
as small rounded outgrowths, two pairs of which belong to each stern-segment. It is noteworthy
that in an early stage of growth (fig. 1) the proximal zooecia are directed towards what may
be called the back of the stem; i. e., the surface opposite to the one on which the brackets of
the distal zooecia are situated. In the older segments all the zooecia may be directed towards
the front of the stem ; but it may be assumed that this position is not constant, and that there
is a considerable range of movement in the fully grown zooecia.
The zooecia are attached by a narrow base, where a joint is present. When fully grown
they are subcylindrical, and are terminated by an orifice, which is quadrangular in shape. The
vestibule is short : — ■ a character which appears to be correlated, in the non-incrusting Ctenostomes
in general, with a feeble development of the collar. This structure is certainly very delicate in
the present species. The tentacles, eight in number, remain unbent during retraction, and the
alimentary canal similarly retains its U-like form (fig. 3). The oesophagus is short. The parietal
muscles are delicate.
The rootlets are well chitinised and may be straight, sinuous or geniculate at intervals;
doubtless in correlation with the nature of the substratum. At the points from which the erect
sterns arise, several internodes may be crowded together. In a particular case (fig. 6) the
arrangement is as follows : — a rootlet (r.1) gives rise to two successive internodes, with no
interval between them. A diaphragm occurs at the commencement of the first internode (e.1),
1) Muscle-fibres of a similar nature have been described by Waters (1910, pp. 239, 250) in Mimosclla graa'/is, Hypophorilla
(fide EHLERS), Farrella rcpais and Valkeria uva. Joi.iet (1877, "Contr. hist. Bry. cötes de France", Arch. Zool. Exp. VI, p. 293) bad
previously described them in Lagenclla nutans (see the following species) and in Valkeria.
80
8i
another between the two internodes, and a third at the distal end of the second internode (z.2),
after which the main rootlet is continued as a delicate thread. The distal internode gives off a
pair of opposite rootlets, which do not branch ; each being separated from the internode by a
diaphragm. The proximal internode similarly gives off two opposite branches, but each of these
at once becomes a short internode (/. z'.1), which gives off two dichotomously arranged branches.
One of these, in each case, continues as an unbranched rootlet, while the other forms a new
internode (/. ï.2), bevond which the branch is continued as a rootlet, while a pair of lateral
rootlets are given off on either side, or one such rootlet on one side. This description may be
summarised by saying that each branching of the rootlets is preceded by the formation of a
short internode, which is separated from each rootlet with which it is in communication by a
diaphragm. Some of these internodes give rise, further, to erect sterns, as shown in fig. 5. On
reversing the slide the base of an erect stem (s) may be seen to originate from the internode i}.
The rootlet marked r? proceeds for some distance without branching, and then forms a new
internode from which another erect stem arises.
The total length of the stem shown in fig. 5, including the part which has been omitted
in the drawing, is 12.35 rnm., or nearly halfan inch. The longest zooecium shown in the same
figure is 750 a in length: while the longer zooecium of fig. 3 is 740 f*, long and 70 u. in greatest
breadth. The proximal stern-segment of fig. 5 is 2 1 60 u. long ; while in the middle of the stem
the segments average about 240 \i. in length and about 40 u. in breadth. The larger rootlets
are 32 0. broad. The diameter of the circular or oval bracket-like supports of the zooecia is
about 20 — 22 u..
The present species is strikingly different from M. gracilis, the genotype of Mimosella.
Some of the most noteworthy respects in which it differs from that species are : — the jointed
character of the erect sterns, the greater relative length of the zooecia and the presence of
four zooecia instead of two on each of the stem-segments.
Mr Waters, who found this species simultaneously with myself, has been kind enough to
suppress the name which he had given to it in favour of the one I had proposed to introducé.
2. Mimosella verticillata Heller. (PI. VII, figs 8 — 10).
Valkeria verticillata Heller, 1867, "Bry. adriat. Meeres", Verh. k. k. zool.-bot. Ges. Wien,
XVII, p. 129, PI. VI, fig. 4.
Hippuraria verticillata Hincks, 1887, uPol. Adriat.", Ann. Mag. Nat. Hist. (5) XIX, p. 311,
PI. IX, fig. 8, Sa.
Hippuraria verticillata Carus, 1889, "Prodr. Faun. Med.", II, p. 51.
Hippuraria verticillata Ostrooumoff, 1896, uOtchet Exp. 'Selianik' "', Buil. Ac. Imp. St-Pétersb.
(5) V, p. 78 (Sea of Marmara).
Lagenella nutans Joliet, 1877, "Contr. hist. Bry. cótes de France", Arch. Zool. Exp. VI,
p. 293, sep. 101 (Roscoff).
Valkeria nutans ) Joliet, 1888, "Ét. Pyrosoma, suivies de Rech. Bry. Roscoff et Menton"
Walkeria nutans S (Paris), p. 106, PI. V, fig. 4.
139. D. Stat. 164. i°42'.5 S„ I30°47.5E., 32 Metres; sand, small stones and shells.
39. C. Stat. Jj. Borneo Bank. 3°27'S., II7°36'E., 59 Metres ; fine grey coral-sand.
(?sp.) 130. F. Stat. 164. i°42'.5 S., I30°47'.5 E., 32 Metres; sand, small stones and shells.
Sr
SIBOGA-F.XPEDIT1E XXVIII ö. I I
82
Stem branching, bearing at intervals groups of sub-verticillate zooecia, arranged on a
series of short internodes given off on each side of a nodal region in the stem. Each of the
lateral internodes bears a pair of elongated, sub-cylindrical zooecia, jointed at the base and
there provided with a series of muscles by vvhich they can no doubt be moved on their stalk-like
proximal end. Vestibule longer than in I\I. óigcwinata, the collar being stronger than in that
species. Tentacles 8, usually bent during retraction ; in which condition the oesophagus is folded
on the pharynx. Gizzard absent.
The structure of the jointed base of the zooecia, with the existence of a cone of muscles,
obviously for moving the zooecium on its stalk, appears to justify the reference of this species
to Mimosella. But the disposition of the zooecia is very different from that found in M. gracilis
and M. bigeminata. The mode of formation of the zooecium-bearing internodes is, however,
not unlike that of the similar structures which, in M. bigeminata, give rise to rootlets; as vvill
be seen by the comparison of fig. 9 with fig. 6. In fig. 9, representing the present species, it
will be seen that the stem gives rise to a swelling (V) cut off distally from its prolongation by
a diaphragm, but not separated from the stem by a proximal diaphragm. On each side, the
swelling in question gives off a short internode vvhich forms the commencement of a series of
similar internodes arranged in such a way as to curve round towards the main stem (see also
fig. 10). Each internode of the lateral series bears a pair of bracket-like projections, similar to
those of M. öigeminata, each of which bears a zooecium. The internode is often prolonged
into an ordinary stem, which after a time may give rise to a new group of zooecia (fig. 10).
In addition to giving off these parts the internode may give off another internode, from one
side; and this in its turn produces another internode, on the corresponding side-, thus giving
rise to the curved series which are so characteristic of this species. A new internode may,
however, be formed medianly, between the two brackets which bear the zooecia.
The zooecium commences with a short cylindrical portion, attached to the bracket by
a joint. It then dilates suddenly, a diaphragm being present in this position (fig. 8). From the
diaphragm originate the fibres of a pair of muscles which together form a cone, their fibres
being inserted into the inner side of the body-wall of the base of the dilated part of the
zooecium. It seems clear, from this arrangement, that there must be some flexibility in the
body-wall at the level of the diaphragm ; and that the two bundies of muscles must be used
for moving the zooecium, as in other species of Mimosella.
The vestibule is of moderate length, and its orifice is square. The collar is sufficiently
strongly developed to be readily seen, in the retracted condition, inside the vestibule (fig. 8). The
tentacles are generally more or less bent during retraction. The oesophagus is of considerable
length and is bent distally on the pharynx when the polypide is retracted. The parietal
muscles form the usual doublé series, extending along most of the length of the zooecium.
They may occur singly, but they are usually in groups of two fibres, and sometimes of a
larger number.
The zooecia are from 700 to 800 u. in length, and usually about 110 to 1 20 u. in breadth.
The zooecium shown in fig. 8 is rather broader than usual, its greatest breadth being 1 70 u..
The main sterns are 30 — 40 n. broad.
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83
The specimen 130. F., although from the same locality as 139. D., appears to belong
to a different species; and I am not even sure that it is a Mimosella. The preparation is
unfortunately not good enough to allow its characters to be described satisfactorily. The zooecia
are shorter than in the other specimens; and their bases do not appear to have the Mintosella-
structure. The mode of branching is similar, though fewer internodes are present. Among the
ordinary zooecia are smaller spindle-shaped structures containing a deeply stained mass, which
appears to consist of a number of longitudinally arranged fibres. I am unable to decide what
is the nature of these structures.
I think there can be no doubt that the specimens 139. D. and 39. C. are very closely
allied to the species described by Heller, Hincks and Joliet, as given in the synonymy at
the commencement of the present account. The balance of evidence seems to be in favour cf
the view that the 'Siboga' specimens belong to Heller's species.
Heller's account was based on material collected on the Adriatic coast, and preserved
in the Triest Museum. The specimens had been labelled by Mexeghixi as Cuscutaria verticillata ;
but this appears to have been merely a MS. name until it was recorded by Heller. The
original account specially mentions the fact that the zooecia commonly originate "paarig (3 — 4
Paare) von einem Puncte"; a description with which my own observations well agree. Hixcks,
some years later, recognised Heller's species from his own Adriatic material. He expressly
comments on the narrow, jointed base of the zooecia : — a feature which had previously been
described by Joliet: — but he describes the "ventral" face of the zooecia as being almost
entirely occupied by a membranous "area"; justifying the reference of the species to Hippuraria
and to the Fam. Triticellidae. If an "area" really occurs in Adriatic specimens the 'Siboga'
form probably does not belong to the same species. I think, however, that it is justifiable to
feel some doubt with regard to the correctness of this part of Hixcks's description ; partly
because Joliet has recorded nothing of the kind in what appears to be the same species; and
partly because no information is given as to the way in which Hixcks' specimens had been
prepared. Unless special precautions are taken in mounting, Ctenostomes of this type become
tnuch shrivelled in the process; and the appearance of a membranous area may have been
due to artificial contraction. But in any case I feel justified, after examining Busk's type-specimen
of Hippuraria egertoni l) preserved in the British Museum in expressing the opinion that Hixcks'
specimens had nothing to do with the form described by Busk. In both the accounts which
have been given by Joliet, that author refers to the narrow peduncle on which the zooecia
execute movements of "nutation". He further describes the muscles by which these movements
are effected and regards them as modified parietal muscles ; and in his later account he describes
their origin from a diaphragm separating the zooecium from its peduncle in a way with which
my own observations agree exactly. There is thus considerable reason for believing that both
the Adriatic specimens and those from Roscoff described by Joliet are referable to Mimosella.
If the 'Siboga' specimens are rightly referred to the same species the distribution of this form
must be a wide one.
1) See below, under Triticella boeckii.
83
84
3. Mimosella tennis n. sp. (PI. VII, figs n — 14).
> Farrella atlantica Waters (nee Busk), 1914, "Mar. Fauna Brit. E. Afr.", "Bry. Cycl ",
p. 852, PI. IV, fig. 9.
? Mimosella sp. Waters, t. cit., p. 852.
' > Stat. 53. Bay of Nangamessi, Sumba, o — 3ÓMetres; Coral-sand ; near the shore, mud.
Stolon very delicate, adnate and branching in a cruciform manner. Zooecia given off
singly, from the middle line, by the internodes, and near their distal ends. The proximal end
of the zooecium forms a narrow, subcylindrical peduncle, of varying length, expanding slightly
at its distal end, where it joins the major part of the zooecium. A joint formed by the partial
ingrowth of a fold of the body-wall is present at this level, and a pair of short muscular fans
dive'rge from the fold and are inserted into the adjacent part of the body-wall of the zooecium,
on the distal side of the joint. The zooecia are elongated and narrow, subcylindrical but tapering
off into the peduncle at the proximal end. Orifice quadrangular, vestibule short. Polypides with
8 tentacles and a short, thick oesophagus. Gizzard wanting.
The present species has been found, in small quantity, from a single Station, where it was
growing on Rctcpora (24. A.), Scritpoeellaria and a Hydroid. It has a considerable superficial
resemblance to Valkeria atlantica, but it is distinguished from that species by the mode of origin
of the zooecia from the stolon and by the joint, with which a pair of muscles are connected,
in the proximal region of the zooecium. This latter character appears to justify the reference
of the present species to Mimosella ; and it is similar to the arrangement which has been
described above in M. verticillata, from which the species differs in the uniserial disposition
of the zooecia, in the greater slenderness of all parts of the zoarium and in the greater length
of the internodes.
The mode of branching of the stolon is represented in fig. 1 1 . The primary internode is
itself somewhat cruciform and is not marked off proximally by a diaphragm. A distal diaphragm
is, however, present; and the internode gives off a short, narrow, tubular portion from its frontal
surface, a short distance on the proximal side of the diaphragm. This tubular portion, which
resembles the similar parts in M. verlicillata, acts as the support of a zooecium. The branches
pass off nearly at right angles from the sides of the internode, and are slender and somewhat
elongated, being terminated distally by a diaphragm, giving off the base of a zooecium in the
same manner as in the primary internode, and sometimes producing lateral branches of a new
order. The peduncle of the zooecium is variable in its length but is usually much longer than
in M. verticillata. The zooecia are relatively much more elongated, in proportion to their width,
than in that species, but otherwise resemble them in structure.
The following are the measurements, in y., of the zooecium shown in fig. 14: — total
length, 1040 ; greatest width, 90 ; length of peduncle, 360. The peduncle of the specimen shown
in fig. 12 is only 250 u. long. The stolon is about 10 p. in diameter.
The unnamed species of Mimosella described by Waters (t. cit.) from Chuaka, Zanzibar,
has considerable resemblances to the present species, but the base of the zooecium is said to
34
be "rounded like that of a Boweröankia". The Farrella atlantica described in the same place
has the muscles of a Mimosella and it is indeed identified by Waters with Hippuraria verti-
cillata Hincks (non Heller). It appears to me to be clearly distinct from F. atlantica Busk.
Fam. Buskiidae Hincks.
Busküdae Hincks, 1SS0, "Hist. Brit. Mar. Pol.", p. 531.
The diagnosis of the Family given by Hincks runs as follows: —
"Zooecia contracted below, not continuous with the creeping stolon, with an aperture
"on the ventral surface".
In the form which I refer doubtfully to Buskia nitens, I have not been able to convince
myself of the existence of the "aperture".
Buskia Alder.
Buskia Alder, 1856, "Cat. Zooph. North. and Durham", Trans. Tyneside Nat. Field Club,
III, Pt 2, p. 156, (sep. 66).
Buskia Alder, 1857, Rep. 26* Meet. Brit. Ass. Cheltenham, 1856, Notices and Abstr., p. 91;
and Quart. J. Mier. Sci., V, "Zoophytology", p. 24.
Buskia liincks, 1880, "Hist. Brit. Mar. Pol.", p. 531.
Buskia Waters, 1910, "Rep. Mar. Biol. Sud. Red Sea", "Bry. II", J. Linn. Soc. Zool., XXXI,
p. 241.
(nee Buskia Reuss, 1865, "Fauna d. deutsch. Oberoligocans, 2. Abth., Sitzb. k. Akad. Wiss., math.
naturw. Cl., L, I Abth., Jahrg. 1864, p. 677). (sep., p. 64).
(nee Buskia Tenison-Woods, 1S77, Papers and Proc. and Rep. Roy. Soc. Tasmania for 1S76, p. 115).
(nee Buskea Heller, 1867, "Bry. adriat. Meeres", Verh. k. k. zool.-bot. Ges., XVII, p. 89).
Mr Waters (t. cit.) states that B. socialis Hincks possesses a gizzard and that it cannot
be included in Buskia (of which the genotype is B. nitens). But if I am right in believing that
both B. nitens and B. setigera possess a gizzard, the subdivision of the genus indicated by
Waters does not appear necessary.
1. Buskia nitens Alder (?). (PI. V, figs 14 — 17).
Buskia nitens Alder, 1856, t. cit., p. 156, PI. VII, figs 1, 2 (sep., p. 66, PI. V, figs 1, 2).
Buskia nitens Alder, 1857, Rep. Brit. Ass. t. cit., p. 91.
Buskia nitens Alder, 1857, Quart. J. Mier. Sci., t. cit., p. 24, PI. XIII, figs 1, 2.
Buskia nitens Hincks, 1880, t. cit., p. 532, text-fig. on p. 533, PI. LXXII, figs 6. 7, ja.
Buskia nitens Hincks, 18S4, "Pol. Q. Charlotte Islands", Ann. Mag. Nat. Hist. (5) XIII, p. 208.
Buskia nitens Levinsen, 1894, "Zool. Dan.", IX, "Mosdyr", p. S3, PI. VIII, figs 12, 13.
Buskia nitens Waters, 1914, "Mar. Faun. Brit. E. Afr.", "Bry. Cycl ", Proc. Zool. Soc,
p. 854.
307. B. Stat. 273. Ofif Pulu Jedan, E. coast of Aru Islands, 13 Metres; sand and shells.
451. C. Stat. 64. Kambaragi Bay, Tanah Djampeah, o— 32 Metres; coral, coral-sand. (On slide
451.B.1, Victorella sibogae).
Zooecia very minute, with an attached base, spinous at its edge, and a short erect
85
86
peristome. Stolon not continuous with the proximal end of the zooecium, very slender, bifurcating
at intervals, the branches usually given off in the interspaces between the zooecia; with a few
transverse diaphragms. Polypide with a gizzard. Collar apparently absent. Tentacles 8.
In its general appearance the present form has a close resemblance to the British species
figured by Aeder and Hincks. The latter author has, however, described an "aperture", or
flat membranous region on the side of the peristome facing the stolon. Alder has, moreover,
described a well developed collar. As I have been unable to find certain evidence of Hincks'
aperture and of Alder's collar, I ara not certain that the reference of the 'Siboga' specimens
to the British species is correct. The species is, however, represented by so small an amount of
material (307. B., one colony; 451. C, two or three zooecia) that I hesitate to describe it as
distinct. It is possible that I have overlooked the parts in question; or that the specimens observed
are to be regarded as a variety in which they are really absent. In view of its minute size it is
not impossible that some of the features ordinarily present are not here developed. Hincks leaves
the existence of a gizzard doubtful; but I have no doubt of its presence in the 'Siboga' material.
It is not easy to be positive with regard to the mode of attachment of the zooecium
to the stolon. At first sight it might be supposed that a narrow, pointed, proximal end of the
zooecium was connected with the stolon. But a more careful examination seems to show that
the pointed end (fig. 16) is merely one of the spines developed from the margin of the zooecium,
the real attachment of which is somewhere near the middle of the adnate portion. This region
is flattened and is provided with a few spinous processes on one side only, instead of on
both sides, as in British specimens. The stolon generally runs along the attached part of the
zooecium, in close contact with it, on the side opposite to that which bears the spines. The
diaphragms of the stolon do not seem to be constant in position.
In one case (fig. 17) the tentacles are protruded, and eight can be counted. In the same
zooecium there is a small emargination on one side of the zooecium, close to the base of the
kamptoderm. This may represent the "aperture"; but I have not been able to find it in other
zooecia. The peristome is usually marked by a number of fine circular striae, and the orifice
is more or less square.
The zooecium shown in fig. 15 is 290 u. long, and 140 u in greatest breadth. The
longest zooecium in fig. 14 is 360 u. long.
Buskia australis Jullien J) seems to be an allied species; but it has a well developed
aperture. No figure is given in the original account. B. socialis Hincks 3) is another form allied
to B. nitens^ from which it is said to differ in its much larger size, as well as by its erect
habit and by other characters.
It might be suggested that this species should be referred to Arachuidiitm, but its mode
of branching seems to exclude this possibility. The cruciform arrangement is not found here,
and the branches of the stolon do not correspond with the bases of the zooecia. It has, however,
some resemblance to Arachnidium simplex, described by Hincks from Barents Sea 3).
1) JUI.I.IEN, J., 1888, p. 22.
2) Hini KS, T., 1887, "Pol. Adriatic", Ann. Mag. Nat. Hist. (5) XIX, p. 310, PI. IX, figs 7-7/'.
3) Ibid., 1SS0, "Hydr. Pol. Barents Sea", Ann. Mag. Xat. Hist., (5) VI, p. 2S4. PI. XV, figs io. II.
S6
87
2. Buskia setigera Hincks. (PI. V, figs S — 10).
Buskia setigera Hincks, 1887, "Pol. Hydr. Mergui Arch.", J. Linn. Soc. Zool., XXI, p. 127,
PI. XII, figs 9-13.
Buskia setigera Kirkpatrick, 1890, "Hydr. Pol. China Sea", Ann. Mag. Nat. Hist. (6), V, p. 17
(Tizard Bank, 27 fathoms).
Buskia setigera Kirkpatrick, 1S90, "Hydr. Pol.Torres Str.'', Proc. R. Dublin Soc. (X. S.) VI, p.612.
Buskia setigera Thornely, 1905, 'Herdmax's "Rep. Pearl Oyster Fish. G. of Manaar", Suppl.
Rep. XXVI, "Polyzoa", Publ. by the Roy. Soc, p. 128.
3S0. F. Stat. 47. Bay of Bima, 55 Metres; mud with patches of fine coral-sand.
322. D. Stat. 50. Bay of Badjo, W. coast of Flores, o — 40 Metres; mud, sand and shells. (On
slide 325. M., Valkeria atlantiea).
339. B. Stat. 71. Makassar, o — 32 Metres; mud, sand with mud, coral (mostly on Hydroids).
38. K. Stat. jj. Borneo Bank, 3°27'S., i\]° 56' E., 59 Metres; fine grey coral-sand.
1 30 N
xf' [ Stat. 164. i°42'.5 S., I30°47'.5 E., 32 Metres; sand, small stones and shells (on
J:>' " l Hydroids, Bugula, etc).
505. D. ] ' o < 1
144. D. Stat. 166. 2°28'.5S., i3i°3'.3E., 118 Metres; hard coarse sand (on a Hydroid : —
Cellepora slides 144. C, 144. C.2).
296. D. Stat. 273. Off Pulu Jedan, E. coast of Aru Islands, 13 Metres; sand and shells (on
Catenicella, slide 296. B.1).
also from Torres Straits (Mus. Zool., Cambridge), A. C. Haddon Coll., Reg. Feb. 24, 1898.
Zooecia relatively large. Stolon well chitinised, forming slight expansions separated by
internodes which vary in length. A diaphragm is developed immediately on the distal side of
each internode, and another on each side. From each of the regions corresponding with a lateral
diaphragm a branch is given off, the two branches being nearly always exactly opposite one
another, and standing at right angles to the main stem. Each lateral branch gives rise to an
internode, which may be formeel close to the parent-stem. Y\ 'hen this is the case in both lateral
branches (fig. 8) the zooecia appear paired, their principal axis being parallel to the main stolon.
In other cases the first node of a lateral branch may be at a greater distance from the main
stolon. The lateral branches do not usually give off new branches; but each is prolonged beyond
the first node (a) as a short barren stolon ; (ö) as a continuation of the branch forming a new
internode which bears a single zooecium and is often prolonged as a barren stolon; (c) as a
stolon which branches like the main stem. The zooecia are almost entirely erect, broader basally
than distally, and with a small attachment to the internode. One aspect of the zooecium is
convex, while the opposite side is fiattened and forms a membranous "aperture". The basal
end of the zooecium gives off a varying number of spines, which may be used to give a firmer
attachment. One of these commonly lies on the stolon, to which it is fastened along its whole
length, in such a way as to form a buttress supporting the base of the zooecium. A pair of
spines, near the proximal end, may be more developed than the others and are directed in an
oblique direction. At its distal end the zooecium bears several spines, in most cases four in
number, arranged at equal distances round the region of the orifice. Each of these spines
consists of a short basal segment foliowed by an elongated, slender, pointed spine. Collar greatly
developed, its supporting ribs at first spirally arranged and then becoming straight (fig. 9).
Polypide with a gizzard, and apparently with 8 tentacles.
87
88
Recorded by Hincks from the Mergui Archipelago (W. of the Malay Peninsula); by
Kirkpatrick from the Tizard Bank (between the Philippine Is and Cochin China) and Torres
Straits, and by Miss Thornely from Ceylon.
Tliis species is obviously a common Ctenostome in the region investigated by the 'Siboga',
as is evident from the list of localities given. It has been well described and figured by Hincks,
to whose account I have little to add. The number of spines at the distal end of the zooecium
is nearly always four, the number recorded by Hincks. But in one case (38. K.) I find a lar°"er
number in some of the zooecia, indicated by their basal segments, which may be as many as
eight. Hincks has given a "correction" (op. cit., foot-note on p. 127) of the position of the
spines shown in tvvo of his figures-, stating that these originate from the margin of the distal
end of the zooecium. The 'Siboga' specimens show, however, that there was no inaccuracy in
the figures in question, but that the spines may appear to originate from the margin or more
proximally, according to the state of protrusion of the kamptoderm.
Hincks has described a peculiar arrangement, which I can confirm, of the supporting
ribs of the collar. These are spirally disposed in the proximal part of the structure (fig. 9),
but become straight distally. By focussing down through the spiral part it can be seen that
the ribs of the opposite side cross those of the side which is uppermost; indicating that the
direction of the spiral is the same all round the collar.
Hincks has stated that that the polypide is "small, and of very simple structure"; and
further that "there seems to be no tracé of a gizzard". I cannot agree with either of these
statements. The polypide is not simpler in its structure than is that of other Ctenostomes. The
vestibule is of great length (fig. 10), in correlation with the great development of the collar;
and it is retracted by strong parieto-vaginal muscles. The tentacles are bent in retraction ; and
the oesophagus in this position passes distally, being, moreover, of considerable length. At its
distal end, this limb of the alimentary canal passes proximally and forms a well marked gizzard
before opening into the stomach. It need hardly be remarked that these relations would be
altered in the protruded condition of the polypide.
The membranous "aperture" varies in length. In the specimen shown in fig. 10 it is
shorter than usual ; and three groups of parietal muscles are visible, on each side; their fibres
being inserted into the membrane of the aperture, the function of which is thus, as might have
been expected, to permit the protrusion of the polypide.
The zooecia are usually from 4S0 u. to 550 ij. long, and as much as 180 \j. in greatest
breadth. The collar shown in fig. 9 is 460 u. long and 1 30 y. broad at its distal end.
The species which has been described by Osburn l) as Hippuraria armata (= Vesicu-
laria armata Verrill, 1874), from the Woods Hole Region, Mass., appears to me to be a
Buskia and to be nearly related to B. sctigera. It may be noted that Osburn describes a
distinct gizzard. It is pointed out below (p. 90) that Busk's original account of Hippuraria
was based on a misconception, and that there is no reason for separating that genus from
Triticclla.
I) OsiiURN, R. C, 1912, "Biy. Woods Hole Region", Buil. Bureau Fisheries (Washington), XXX, p. 256, PI. XXIX, figs S4, 84^.
88
§9
3. Buskia pilosa n. sp. (PI. V, figs 11 — 13).
Type. 501. E. Stat. 163. Near Seget, W. entrance Selee (Galewo) Strait, 29Metres; sand and stone,
mixed with mud (on rootlets of Caberea lata).
Also from Torres Straits (Mus. Zool., Cambridge), A. C. Haddon Coll., Reg. Feb. 24, 1898
(on a Hydroid).
Zooecia very large, more or less plano-convex in shape, attached by a flattenecl base,
which is generally at least two thirds of the entire length of the zooecium. The basal wall is
thin and membranous, and is continued along one surface of the free portion as far as the
orifice. Frontal and lateral surfaces very convex, covered with a thick cuticle, all parts of which
give off a profusion of spines, which are of several kinds. Stolon slender, very irregular and
frequently geniculate; dilating at intervals into slight expansions, from which branches are given
off, as in B. setigera.
The zooecia are all dead and empty ; and there is thus no certain proof that the organism
here described is a Polyzoon. But I think there can be no doubt of this-, and the species has
sufficiënt resemblance to B. setigera to justify its reference to the same genus. The cuticle of
the free surface is very thick and stout; since in the Torres Straits specimen, which is a dry
mount, the zooecia show no noticeable shrinkage, and agree precisely in form with the 'Siboga'
specimens mounted in glycerine or Canada balsam. The continuation of the flat, membranous
base as far as the orifice indicates that an aperture is represented, as in typical species of
Buskia. The spines are a specially characteristic feature of this form, and the following kinds
may be distinguished : — (a) those of the greater part of the free surface, which are relatively
short, pointed and curved ; (ó) a tuft of much longer spines, which appear to correspond with
the oral spines of B. setigera : they are, however, much more numerous than in that species
and are not jointed; while they usually have a sinuous outline; (e) those of the edge of the
convex portion of the zooecium, which are commonly bifurcate and are used for attachment to
the object on which the colony is growing.
The distal region of the zooecium is free, and forms a short "peristome", the orifice
being no doubt terminal and surrounded by the oral spines ; though the orifice cannot actually
be demonstrated except so far as it is indicated in fig. 13. Xone of the details of the internal
anatomy can be made out ; as the zooecia are quite empty, with the exception of one or two
which contain a multinucleated mass which looks like an encysted (intrusive) Protozoon.
The stolon has a close resemblance to that of B. setigera ; the swellings from which
branches are given off being much like those of that species. But the stolon of the present
species is very irregular in outline and is frequently angulated. It is of considerably smaller
diameter than the Gz^r^tf -rootlets (fig. ii,r) over which it creeps.
The zooecium shown in fig. 13 measures 600;/ in length without its spines: and is
260 u. in greatest diameter, without the spines. The finer stolon-threads are 10 [i in diameter,
while the width of the Caócrea-rootlets is about 50 ft.
S9
SIBOGA-EXPED1T1E XXVIII».
9o
Fam. Triticellidae G. O. Sars.
Tritilellidae (misprint for Triticellidae), Sars, 1874, "Om en . . maerk. Slaegtstype af Polyzoer",
Forh. Vid.-Selsk. Christiania, Aar 1873, p. 397 (sep., p. 13).
Triticellidae Hincks, 1880, "Fust. Brit. Mar. Pol.", p. 541.
This Family is not represented in the collection here reported on; but as the distribution
of Triticella boeckii noted below gives sorae reason for supposing that it is not unlikely to
occur in the Malay region, a few notes on this species may not be out of place.
Triticella Dalyell.
Triticella Dalyell, 1848, "Rare and remark. An. Scotland", II, p. 66.
Cordyle Boeck, 1862, "Forelob. Bemaerkn. Pol.", Forh. Vid.-Selsk. Christiania f. 1861, p. 49.
Triticella G. O. Sars, 1874, 1. cit.
Hippuraria Busk, 1874, "Not. New Pol.", Proc. Zool. Soc., p. 29.
Triticella Hincks, 1880, t. cit., p. 542.
Triticella Duerden 1893, "New and rare Irish Pol.", Proc. R. Irish Acad. (3) III, p. 129.
1 . Triticella boeckii G. O. Sars.
Triticella boeckii Sars, 1874, t. cit., pp. 387, 397 (sep., 3, 13), PI. VIII.
Triticella boeckii Carus, 1889, "Prodr. Faun. Med.", II, p. 51 [Mediterranean].
Triticella boeckii Duerden, 1893, t. cit., p. 131.
Triticella boeckii Duerden, 1895, "Surv. Fishing Grounds, W. Coast Ireland", "Hydr. and Pol",
Proc. R. Dublin Soc. (N. S.) VIII, p. 334-
Triticella boeckii Ostrooumoff, 1896, "Otchet Exp. 'Selianik'", Buil. Ac. Imp. St. Pétersb. (5)
V, pp. 37, 66 (on Geryon tridens, Sea of Marmara).
Hippuraria egertoni Busk, 1874, 1. c.
The genotype of Triticella is T. jïava Dalyell. A species of this genus had previously
been recorded by Gros1), under the name of Phimatella familiaris, from Ostende and Havre;
and this specific name has been adopted by Smitt 2), for one of the species of the genus,
apparently identical with T. korenii Sars.
Mr Waters8) has made the following statement: - - "Hippuraria, Busk, has a delicate
"rhizome, from which groups of zooecia arise. In the only specimen (which is now in the British
"Museum) it is growing upon the stalk of a seaweed, and this was mistaken for the stem of
"■Hippuraria. There is, therefore, now absolutely no reason for separating Hippuraria from
" Triticella'.
I agree with Mr Waters in his general conclusion, but from an examination of the type-
specimen (73. 11. 21. 1) of H. egertoni, to which he alludes, I have come to a different conclusion
with regard to certain details. The slide had suffered from the drying up of the medium in
which the specimen had been mounted; and it has been re-mounted by Mr Kirkpatrick since
Mr Waters examined it. It is now quite clear that the description "delicate rhizome" is
i) Gros, G., 1849, "Fragm. d'Helminthologie", Buil. Soc. Imp. Nat. Moscou, XXII, p. 567, PI. VI, figs G, 1 — 10.
2) Smitt, F. A., 1866, "Krit. fort. Skand. Hafs-Bry.", II, Öfv. K. Vet.-Ak. Förh., 1S66, pp. 502, 524, PI. XIII, fig. 36.
3) Waters, A. W., 1910, p. 241, note.
90
9i
unsuitable. The specimen consists of several groups of zooecia, each of which arises from an
isolated base completely surrounding the jointed tubular object which Busk mistook for the stem
of the Hippuraria. The base is of precisely the same nature as that described as a "continuous
crust" by Duerden (1893), in Triticella boeckii and other species. The zooecia show the gibbous
"dorsal" outline which is so characteristic of that species; and agree with it in the position and
general development of the curved chitinous thickening of the ectocyst described by Duerden as
the "frenaculum". The comparison of Busk's specimen of Hippuraria egertoni with undoubted
specimens of T. boeckii leads me, in fact, to the conclusion that the two forms are identical.
The label of Busk's slide states that the specimens was "Attached to eye of Gonoplax
angulatus' . An examination of the supposed stem of the Polyzoon shows that it is a jointed
structure, bearing rings of minute spines at the joints. Mr Waters regarded it as a seaweed,
but it appears to me nearly certain that it is really a fragment of the flagellum of the antenna
of the Gonoplax. Dr W. T. Calman, whose opinion I asked, agrees with this conclusion. The
base of the antenna lies close to that of the eye in Gonoplax, as may readily be seen by
examining a specimen of the crab. When it is remembered that Busk mistook the antenna for
part of the Hippuraria, the statement that the Polyzoon was attached to the eye of the crab
becomes readily intelligible.
In order to obtain further evidence with regard to the nature of Hippitraria I asked
Dr Calman to select specimens of Gonoplax which seemed likely to bear this interesting form.
He was kind enough to send me specimens, from the collection in the British Museum, on
several of which there was 110 difhculty in finding perfectly typical specimens of Triticella
boeckii. All the crabs are labelled Gonoplax rhomboides Linn., with which, as Dr Calman informs
me, G. angulata Fabr. is generally regarded as synonymous. The specimens on which T. boeckii
was found are as follows : —
(1) 98.5.7.251 — 252. Bay of Biscay, "Porcupine", 1870, Coll. Norman ; on 5 specimens, attached
principally to the bases of the walking legs, particularly on the dorsal side, in the angles
between the legs and the carapace.
(2) 1 9 1 1 . 11. 8. 34. Off Valentia, Ireland, "Porcupine", 1869, Stat. 6, 90 faths, Coll. Norman;
on 1 specimen.
(3) 191 1. 1 1. 8. 3 1. Cadiz Harbour, Coll. Norman ; on 1 specimen.
(4) 191 1. 11. 8. 35 — 38. Off Cape Sagres [Portugal], 45 faths, "Porcupine", 1S70, Coll. Norman;
on 1 specimen.
(5) 1910. 2.4. 180 — 182. Same locality; on 1 specimen.
(6) 96. 5. 195 — -198. Algoa Bay, H. A. Spencer; on 4 specimens, growing on the bases of the
legs and great chelae; also on the third maxillipedes, and on the edges of the abdomen.
(7) Crab not determined [Gonoplax sp.), 16 miles N. E. of Bird Is [Algoa Bay], 40 faths,
F. Toppin 1905; on 1 specimen, growing on the bases of the walking legs; on the second
and third maxillipedes; on the "front", close to the antennae; in the orbits; on the back
of the carapace; and on the abdomen.
The localities given for T. boeckii by Duerden (1893) are Berehaven, S. W. Ireland
91
92
(which, it may be noted, is the locality from which Busk's Hippuraria egertoni was obtained),
on Portunus depmrator\ and Sars' record from Christiania, on Geryon tridens. In 1895 Duerden
mentions the occurrence of the same species, from another Irish locality, on Gonoplax angulata,
in special abundance on the anten na e. The list which I have given above shows
that it is of common occurrence in more southern localities; and in particular on Gonoplax
from Portugal, Spain and Algoa Bay. The last locality suggests that an examination of the
Crabs and other Crustacea from the Malay region might be expected to result in the discovery
of specimens of Triticella. It may be noted that Annandale x) has recorded T. korenii from
Squilla fasciata obtained off Japan.
If Hippuraria is a synonym of Triticella, it follows that the species placed in the former
genus by previous authors must find a place elsewhere. I have cited H. verticillata Hincks, and
II. armata Osburn under Mimosella verticillata and Buskia setigera respectively. H. elotigata
Osburn (t. cit., p. 256) appears to be a Triticella.
D i s t r i b u t i o n of Entoprocta and Ctenostomata.
STATION
DEPTH in Metres
ENTOPROCTA
CTENOSTOMATA
43
47
50
53
58
60, 303
64
71
0—15
0—36
55
o — 40
0—36
o — 27
0—36
0—32
Barentsia gracilis.
Loxosoma circulare (Retepora).
Barentsia taxa.
Loxosoma troglodytes (Lepralia).
Barentsia gracilis.
Loxosoma circulare (Retepora).
Loxosoma velatum (Retepora).
Loxosoma annulatum (Retepora).
Barentsia gracilis.
Loxosoma loricatum (Canda).
Barentsia gracilis.
Loxocalyx lineatus (Halichondria I.
Loxosoma cirriferum ( Retepora 1.
Barentsia gracilis.
Barentsia discreta.
Bowerbankia imbricata (?)
Mimosella bigeminata.
Amathia distans.
Valkcria atlantica.
Nolella annectens.
Valkcria atlantica.
Buskia setigera.
Nolella papuensis.
i Imathia distans.
Boiverbankia imbricata (
Valkcria atlantica.
Buskia setigera.
Mimosella tennis.
Amathia distans.
Victorella sibogae.
Bowerbankia imbricata (?)
Valkcria tuberosa.
Buskia nitens.
Victorella sibogae.
Nolella papuensis.
i Iracknoidea protecta.
Bowerbankia imbricata (?)
1) Annandale, N., 1912, "Pol. Indo-Pacific Stomatopods", Ree. Ind. Mus. VII, p.
92
[24.
93
STATION*
DEPTH in Metres
ENTOPROCTA
CTEXOSTOMATA
71
77
80
91
0—54
99
16—23
105
275
"5
Reef
116
72
117
80
133
0—36
144
156
162
163
164
0—^2
213
240
250
257
273
59
40—50
45
469
iS
29
32
166
118
81 & 231
36—54
204
75—94
-36
9—45
20—45
o — 52
Loxosoma sluiteri.
Barentsia laxa.
Loxosoma sp. (Retepora).
Loxosoma loricatum (Canda).
Loxosoma pusilhim (Retepora).
Loxosoma sp. (Retepora).
Barentsia gracilis.
Loxosoma cocciforme (Siplionicytara).
Loxosoma annulatum (Retepora).
Loxosoma pusillum (Retepora).
Loxosoma breve (Schisoporella).
Loxosoma troglodytes (Lepralia).
Barentsia discrcta.
Loxocalyx leptoclini (Ceplialodisats).
Loxosoma sp. (Retepora).
Barentsia gracilis.
Barentsia discreta.
Loxosoma circulareQ) (Retepora).
Barentsia gracilis.
Loxosoma loricatum (Canda).
Loxosoma loricatum (Canda).
Loxosoma cirriferum (Retepora).
Pedicellina compacta.
Barentsia gracilis.
Valkcria atlantica.
Buskia setigera.
Mimosella bigeminata.
Mimosella verticillata.
Buskia setigera.
Nolella papuensis.
Valkeria tuberosa.
Nolella papuensis.
Amathia convoluta (?).
Nolella papuensis.
Valkeria atlantica.
Valkeria atlantica.
Valkeria tuberosa.
Nolella papuensis.
Valkeria atlantica.
Nolella papuensis.
Valkeria atlantica.
Mimosella bigeminata.
Valkeria atlantica.
Buskia pilosa.
Nolella papuensis.
Amathia convoluta.
Bowerbankia imbricata (?).
Valkeria atlantica.
Valkeria tuberosa.
Mimosella bigeminata.
Mimosella verticillata.
Mimosella (? gen. & sp.).
Buskia setigera.
Buskia setigera.
I 'alkeria atlantica.
Nolella papuensis.
Arachnidiu m irregulare.
Nolella papuensis.
Valkeria atlantica.
Valkeria tuberosa.
J Iimosella bigem inata.
Nolella papuensis.
Vesicularia papuensis.
Amathia convoluta.
Amathia distans.
Valkeria atlantica.
Buskia nitens.
Buskia setigera.
93
94
STATION
DEPTH in Metres
ENTOPROCTA
CTENOSTOMATA
-74
57
Loxosoma annulatum ( Retepora).
Nolella papuensis.
Loxosoma cirriferum (Retepora).
Amathia convoluta.
Loxosoma pusillum (Retepora).
Barentsia discrela.
282
27—54
Loxosoma lor i ca turn (Canda).
2S5
34
Loxosoma sp. (Retepora).
305
113
Loxosoma sp. (Retepora).
Loxosoma loricatum (Canda).
310
73
Loxosoma annulatum (Retepora).
3"
0—36
Valkeria tuberosa.
318
88
Loxosoma subsessile (Conescharellina).
Barentsia gracilis.
Valkeria atlantica.
319
82
Barentsia gracilis.
Barentsia geniculata.
320
82
Barentsia gracilis.
321
82
Loxosoma annulatum (Schizoporella).
The preceding list gives a complete statement of all the specimens of Entoprocta and
Ctenostomata which I have found among the 'Siboga' dredgings. It may be remarked, in the
first place, that the conditions which are favourable for the occurrence of either of these groups
appear to be favourable for that of the other; — since Entoprocta and Ctenostomata were
found in company in a considerable number (17) of the Stations. Only 42 of the Stations have
yielded both groups or one of them, out of a total of about 135 Stations at which Polyzoa were
obtained. Entoprocta are represented at 30 Stations, and Ctenostomata at 29 Stations.
So far as the Loxosomatidae are concerned, the distribution is no doubt affected by that
of the animal on which they are found. Loxosoma cocciforme was discovered in the material
from Stat. 156, at a depth of 469 Metres. I believe this to be the greatest depth from which
any species of this genus has been obtained; but it seems natural to suppose that this exceptional
depth depends largely on the fact that Siphonicytara, the Cheilostome Polyzoon on which the
Loxosoma was found, is in some way adapted for an abyssal existence. Loxosoma sluiteri, on
Phascolion, was obtained from Stat. 105, 275 Metres. With these exceptions, only one of the
above Stations exceeds 100 Metres; namely Stat. 166 (118 Metres), at which a single Ctenostome,
Buskia setigera, was found. Nolella papuensis and two species of Loxosomatidae occurred at
Stat. 204 (75- — 94 Metres); while Valkeria atlantica is recorded from Stat. 117 (80 Metres) and,
in company with two Entoprocta, from Stat. 318 (88 Metres). All the rest of the Ctenostomata
were found at depths of less than 60 Metres, and most of them in still shallower water.
The majority of the Entoprocta similarly come from dejiths of less than 60 Metres.
Those from greater depths, in addition to the records indicated in the preceding paragraph,
are: - ■ Stations 319, 320 (82 Metres), two species of Barentsia-, Stat. 318 (88 Metres) one
Barentsia, and one Loxosoma on a Cheilostome (Couesc/iarellina). The remaining records
exceeding 60 Metres consist entirely of species of Loxosoma growing on Cheilostomes, and are
as follows: Stat. 116 (72 Metres), one species, on Retepora; Stat. 305 (113 Metres), two
94
95
species, on Retepora and Canda respectively ; Stat. 310 [ji Metres), one species, on Retepora\
and Stat. 321 (82 Metres), one species, on Schizoporella.
It is thus clear that the Entoprocta and Ctenostomata of the Malay Region, as in other
parts of the world, are in the main shallow water forms. Of the former group, with the exception
of two abyssal species of Loxosoma (469, 275 Metres) and of others from 73 to 113 Metres,
whose distribution may be considered to be affected by that of their "host"', Barenlsia seems to
descend to a slightly greater depth than the Loxosomatidae.
The distribution of Entoprocta and Ctenostomata in the Archipelago, as revealed by
these records, is curiously unequal. Thus from the whole series of Stations (93 — 109) from the
Sulu Sea, the only member of either group was Amathia convoluta (? sp.), from Stat. 99 (16—
23 Metres), although this particular Station was rich in Polyzoa generally, and was, moreover,
well within the limits of depth at which the two groups were found to flourish in other regions.
Ctenostomata were obtained principally in two well marked groups of Stations :
(I) off the N. W. extremity of New Guinea, including the region from the Aru Islands to the
S. coast of Ceram, and the S. point of Halmaheira (Djilolo); (II) off the chain of Islands
stretching from Java to Flores, in the Straits of Makassar and on the other coasts of Celebes,
and just extending to the Sulu Sea. Only one Station (133, Talaut Is) at which Ctenostomes
were found falls outside these limits. The map published by Prof. Max Weber in his preliminary
account v) of the Expedition shows that the two principal areas correspond with two relatively
shallow water plateaux separated by deep water. One of these extends from the N. coast of
Australia to New Guinea, and the other from Borneo through Java and other Islands to Flores,
and, in other directions, to Celebes and the Sulu Sea. It is noteworthy that the coast of Timor,
which is separated from Flores by deep water, has yielded no Ctenostomes, though Elzerina
blainvillii was recorded by Lamocroux from Timor.
The individual Stations from which Ctenostomes were obtained in greatest abundance
were: — from area (I), Stat. 164, W. of the N. end of Xew Guinea (9 species) and Stat. 273,
Aru Is (6 species): — and from area (II), Stat. 50, W. end of Flores (5 species); Stat. 64,
Tanah Djampeah, X. of Flores (4 species); and Stat. 71, Makassar (6 species). The depth of
none of these dredgings exceeded 40 Metres. At all these Stations a considerable number of
other Polyzoa were obtained, Stat. 164 being specially rich in these animals. It has been pointed
out by Prof. Weber2) that Stations 164 and 273 were characterised by the richness of their
fauna in general.
A careful tabulation of the occurrences of the individual species has failed to reveal
anything that could be held to justify the division of the entire area covered by the 'Siboga'
dredgings into sub-regions. Omitting the two species [Alcyonidium polyoum and Elzerina
blainvillii) which were obtained from Torres Straits, outside the 'Siboga' area, 18 species of
Ctenostomata are described in the present Report. Of these, 14 occur in the first of the large
areas indicated above, and 13 in the other area. The two species found in the rather isolated
1) Weber, M.. 1900, uDie Niederlünd. 'Siboga' Exped.". Petermann's Geogr. Mitth.. Heft VIII. p. 3 (sep.).
2) Weber, M.. 1902, Monogr. I, "Introd. et Descr. de 1'Expéd., pp. 72, 124 — 126.
95
96
Station 133 occur in both the large areas. The species which were obtained most frequently
are scattered throughout the region from which Ctenostomes were obtained. This is the case,
tor instance, with Valkeria atlantica, which was found at 14 of the 29 Stations at which the
group was represented; while Nolella papuensis occurred at 12 Stations, Buskia setigera at 7,
and Valkeria tuöerosa at 6. It may be noted, however, that Arachnidiiuii irregulare, Arach-
noidea protecta and Victorella sibogae were only found at Stations 64, 71 and 213, three
localities lying close together off the S. end of Celebes. But as the first two of these species
were each obtained at only one Station, and the third at only two Stations, the number of
records is insufficiënt to prove much.
There is perhaps sufficiënt evidence to warrant the general conclusion that Ctenostomes
do not fiourish where they are exposed to the full force of the open ocean. The headquarters
of this group in the Archipelago seem to be, as has already been pointed out, round the
W. end of New Guinea and from Celebes to the Islands from Java to Flores. Their absence
from the more exposed S. coast of Timor, and the fact that there is only one record of their
occurrence in the Sulu Sea ; — in both of which localities a considerable number of dredgings
were made; — may be significant. It is probable that the clistribution of the group depends
more on the nature of the locality (depth, character of bottom, etc.) than on any Geographical
considerations which are perceptible in the area investigated.
It has been pointed out above that the distribution of the Entoprocta conforms in general
to that of the Ctenostomata. It may be remarked, however, that three species of this group
were obtained off the coast of Timor (Stations 60, 282, 285) and that none were found in
the Sulu Sea.
Sub-Order II. CYCLOSTOMATA Busk.
Cyclostomata Busk, 1852, J. MacGillivray's "Voy. 'Rattlesnake'", I, p. 346.
Cyclostomata Busk, 1859, "Monogr. Foss. Pol. Crag", Palaeont. Soc, p. 9.
Fam. Crisiidae Johnston.
Crisiadae (pars), Johnston, 1838, "Hist. Brit. Zooph.", Ed. 1, p. 260.
Crisiadae Johnston, 1847, Ibid., Ed. 2, p. 282.
Crisia Lamx.
Crisia (pars), Lamouroux, 1812, "Mém. class. Pol. coralligènes", Nouv. Buil. Sci. Soc. Philomat.,
III, p. 183.
Crisia (pars), Lamouroux, 18 16, "Hist. Pol. Cor. Flex." p. 136.
Crisia (pars), Lamouroux, 1821, "Exp. Méthod.", p. 6.
Crisia (pars), Lamouroux, 1S24, "Encycl. Méthod.", "Zoophytes", p. 224.
1. Crisia elongata Milne Edwards. (PI VIII, figs 1 — 8).
Crisia elongata Milne Edwards, 1838, "Mém. Crisies", Ann. Sci. Nat., Zool., (2) IX, p. 203,
PI. VII, fig. 2 (Red Sea).
96
97
Crisia élongata Busk, 1875, "Cat. Mar. Pol. Brit. Mus.", III, p. 5, PI. IV, figs 5, 6 (Algoa Bay).
Crisia élongata Waters. 1S79, "Bry. Naples", Ann. Mag. Nat. Hist. (5) III, p. 269, PI. XXIII,
fig. 1 (Naples, 40 fathoms).
Crisia élongata Busk, 1S86, Challenger Rep., Pt L, p. 5, PI. I, figs 3, 3*2 (\V. of Fiji Islands,
1450 fathoms).
Crisia élongata Kirkpatrick, 1890, "Hydr. Pol. China Sea", Ann. Mag. Nat. Hist. (6) V, p. 17
(Tizard Bank, 27 fathoms).
Crisia élongata Waters, 1914, "Mar. Faun. Brit. E. Afr.", "Bry. Cyclost.", Proc. Zool. Soc,
p. 838, PI. I, figs 3, 4; PI. IV, fig. 6.
? Crisia sinensis D'Orbigny, 1850 — 1S52, "Pal. Franc. Terr. Crét."', V, p. 599 (China Sea, recent).
Mr Waters (1905, "Recent Bry. D'Orbigny's Coll." Ann. Mag. Nat. Hist. (7) XV, p. 13)
was unable to refer the fragments in D'Orbigny's Collection to any other species.
? Crisia punctij 'era Hasvvell, 1879, "Cycl. Pol. Port Jackson", Proc. Linn. Soc. N. S. Wales,
IV, p. 355.
Crisia terrae-reginae, Hasvvell, 1880, "Pol. Queensland", Proc. Linn. Soc. N. S. Wales, V,
p. 35, PI. I, fig. 1.
Crisia nigrijuncta Ortmann, 1889, "Japan. Bry.", Arch. f. Naturg. LVI, I, p. 58, PI. IV,
fig. 19.
Crisia denticnlata Philipps, 1899, WlLLEY's "Zool. Res.", Pt IV, pp. 441, 449 (Lifu, Loyalty
Islands).
Crisia denticnlata Waters, 1910, "Rep. Mar. Biol. Sud. Red Sea", "Bry. II", J. Linn. Soc, Zool.,
XXXI, p. 232, PI. XXIV, figs 1 — 3, PI. XXV, fig. 11.
? Crisia denticnlata var. gracilis Busk, 18S6, Challenger Rep., Pt L, p. 5, PI. I, figs 4 — \d
(Philippine Islands).
Crisia acropora MacGillivray, 1895 (nee Busk), "Monogr. Tert. Pol. Vict.", Trans. R. Soc.
Vict., IV, p. 118, PI. XVI, figs 2, 8, 9. (The characters of the internode and the form of
the ovicell make this identification probable.)
Crisia holdswort/ui Thornely, 1905 (nee Busk), Herdman's "Rep. Pearl Oyster Fisheries
G. of Manaar", Suppl. Rep. XXVI, Publ. by the Roy. Soc, p. 127.
? Crisia margaritacea Busk, 1875, "Cat. Mar. Pol. Brit. Mus.", III, p. 6, PI. VI. B, fig. 1 (Australiai.
427. C. J_ \. Haingsisi, Samau Island, Timor, o — 36 Metres; Lithothamnion.
( Stat. 303 )
(?sp.) 37. R )
. TT ( Stat. "j-j. Borneo Bank, 3°27'S., ii7°36'E., 59 Metres; fine grey coral-sand.
57. L. Stat. 80. Borneo Bank, 2°25'S., U7°43'E., 40 — 50 Metres; fine coral-sand.
459. G. Stat. 81. Pulu Sebangkatan, Borneo Bank, 34 Metres; coral-bottom and Lithothamnion.
475. A. Stat. 105. 6°8'N., i2i°i9'E., 275 Metres; coral-bottom.
101. B. Stat. 133. Anchorage off Lirung, Salibabu Island, o — 36 Metres; mud and hard sand.
107. B. Stat. 144. Anchorage N. of Salomakiëe (Damar) Island, 45 Metres; coral-bottom and
Lithothamnion.
117. B. Stat. 164. i°42'.5 S., I30°47'.5 E., 32 Metres; sand, small stones and shells.
343. A. Stat. 213. Saleyer, o — 36 Metres; coral-reefs, mud, and mud with sand.
350. B. Stat. 240. Banda anchorage, 9 — 45 Metres; black sand, coral.
5*41. C. Stat. 257. Du-roa Strait, Kei Islands, o — 52 Metres; coral.
3 T 2 A )
' ' | Stat. 273. Anchorage off Pulu Jedan, Aru Islands, 13 Metres; sand and shells.
544. B.)
194. C. Stat. 274. 5°2S'.2S., I34°53'.9E., 57 Metres; sand and shells, stones.
208. B. Stat. 282. Anchorage between Nusa Besi and the N. E. point of Timor, 27 — 54 Metres;
sand, coral and Lithothamnion.
' Stat. 310. 8° 30' S., U9°7'.5E., 73 Metres; sand with a few pieces of dead coral.
253. A. Stat. 315, Anchorage E. of Sailus Besar, Paternoster Islands, o — 36 Metres; coral and
Lithothamnion.
97
SIBOGA-EXPEDITIE XXVIII il. 17.
98
I refer the following specimens, in the collection of the University Museum of Zoology,
Cambridge, to the present species : —
Torres Straits, A. C. HADDON ColL, Reg. Feb. 24, 189S (Slides 10, 26, 96, 158).
Ceylon, Miss L. R. TlIORNELY, Reg. Apr. 25, 1906, 70 (determined by Miss THORNELY as
C. holdsworthii).
Port Denison, Queensland, Miss E. C. Jelly, Reg. May 24, 1895 (3 slides).
Port Denison, Queensland, Australian Museum, Sydney, Reg. Oct. 23, 1899.
Lifu, Loyalty Islands, A. WlLLEY Coll., Reg. Mar. 1, 1898 (determined by Miss PHILIPPS as
C. denticulata).
Japan, found among rooting spicules of Enplectella marshalli, Prof. K. MiTSUKURl, Reg.
Sept. 23, 1896.
Red Sea, T. HlNCKS Coll., Reg. May 13, 1899 (determined by Mr HlNCKS as C. clongata).
Off Fiji, Stat. 176, Challenger Coll., 1450 fathoms, Reg. Nov. 18, 1899.
Zoarium relatively large, well branched. Internodes narrow in the proximal parts of the
colony; becoming broad more distally, and then commonly with a sigmoid fiexure. The frontal
surface of the internode has a broad, slightly convex region in which the outlines of the zooecia
are hardly visible. At the edges of the branch, the individual zooecia become distinct, terminating
in a short peristome which is abruptly bent frontally, the lateral outer margin of the zooecium
being often slightly carinate, the keel occasionally produced at its distal end into a point which
occurs at the base of the peristome. The number of zooecia in an internode may be even or
odd and is extremely variable ; in elongated internodes reaching a high figure. Branches given
off moderately high in the internode, seldom if ever from the lowest zooecium, usually not
lower than the third of one side. Basis ra mi more or less wedged in between two zooecia.
An ordinary internode may give off either one or two branches. A fertile internode typically
bears the ovicell near the middle of its length and gives off three branches, one on the proximal
side of the ovicell, the other two just distally to it. Ovicells with very numerous pores; greatly
dilated distally, the most prominent part being near the distal end. Ooeciostome l) not developed.
Ooeciopore ]) a transverse slit in the distal wall of the ovicell, close to the point where it joins
the frontal surface of the branch. Spines wanting. Joints black. Rootlets arising principally from
the proximal internodes, black-jointed ; occasionally terminating in a curved segment of the form
shown in fig. 2 -).
The present species belongs to a group characterised by the considerable breadth of the
internodes; a feature which is due to the fact that the proximal ends of the zooecia extend
along the median sides of their predecessors, thus giving rise to the appearance of a broad
region, in the middle of the frontal side of the branch, between the distal parts of the preceding
zooecia. A number of species with the same general appearance have been described as Tertiary
fossils, as for instance C. h'órnesii Reuss 3) and C. s?/óaet/ua/is Reuss 4). The fossil forms occur
1) Ooeciostome = the tube by which the larvae escape from the ovicell; ooeciopore = its external orifice (Harmer, 1S9S,
"Dev. Tuiuiipora", Quart. J. Mier. Sci., XLI, p. 81).
2) Rootlets of this type have been described by Busk (1875, p. 6) in C. acropora\ and also, if I understand the description
correctly, by Levinsen (1912, "Stud. Cycl. Operculata", Kgl. Danske Selsk. Skr. (7) Naturv. og Math. Afd., X, 1, p. 16, note), ia a species
mentioned, but not described, as C. hamifera n. sp., found at some distance to the West of the Southern part of the L'nited States.
3) Reuss, A. E., 1847, "Koss. Polyparien Wiener Tertiarbeckens", Haidinger's Naturvv. Abhandl., II, p. 54, PI. VII, fig. 21.
4) Ibid., 1869, "Pal. Stud. alt. Tertiarschichten Alpen", II, "Foss. Anth. u. Bry. Crosara", Denkschr. k. Akad. Wiss. Wien,
Math.-Naturw. Cl., XXIX, 1 Abth., p. 279, PI. XXXIV, fig. 8.
98
99
as isolated internodes, in which ovicells are seldom present ; and it seems to me unsafe, in
most cases, to attempt to identify them with recent species. It may, hovvever, be noted that
C. elongata has been recorded in the fossil condition v).
Among recent species the present tbrm has considerable resemblance, in its broad inter-
nodes, its general mode of branching, and its black joints to the well known C. denticulala,
from which it appears to differ in the more inflated shape of its ovicell, in the position of this
structure near the middle of the length of the internode and in the greater profuseness with
which branches are developed in the fertile internode.
Miss A. Robertson has called attention to the importance of the mode of branching
in the fertile internodes '-) ; and this seems to be really a character which can be used with
advantage in the discrimination of the species. These internodes may be regarded as the parts
of the colony in which the definitive characters of the species find their fullest expression ; the
more proximal parts of the zoarium frequently showing characters which are nearly uniform in
several distinct species.
The erect sterns of a Crisia colonv, like those of some of the more delicate branching
Cheilostomes to which I shall have occasion to refer in the later parts of this Report, commonly
arise as branches given off by adnate rootlets 3). The present species has been found to furnish
definite evidence in favour of this view. In its basal region the internodes of the erect branch
consist of a very small number of zooecia, and have at first a diameter which is little greater
than that of the rootlet. In tracing the branch upwards the internodes are found to become
both broader and longer; the number of their zooecia increasing to a corresponding degree.
It is only when growth has gone on for some time that the definitive characters of the species
become marked. It thus follows that young branches, or the proximal parts of older ones, have
an appearance entirely different from that of fully grown specimens ; and in the absence of
other evidence might easily be referred to different species (cf. figs 3, 5). This is of course
true of the genus and not merely of the present species. As I have pointed out in an earlier
paper *), the presence of ovicells is highly desirable, if not essential, in order to arrive at a
definite conclusion with regard to the determination of a specimen. It is fortunate that ovicells
are well developed in much of the 'Siboga' material.
The axis of a branch is usually sinuous, the convexities following one another in regular
alternate order, and each one typically giving rise to a new branch. The branches, like the
convexities of the stem, are thus alternately arranged. The form of the internode depends to
a large extent on the frequency with which transverse joints are developed. This is a very
variable character; and it accordingly follows that the internodes themselves are also variable
in length and in the extent to which they produce branches.
Fig. 1 represents a portion of a colony in which the specific characters are fully developed.
Joints are not numerous and the internodes are thus very long. The longer internodes have
1) Neviani, A., 1891, "Contr. Conosc. Bri. Foss. Ital.", "Bri. postpliocenici Livorno'', Boll. soc. geol. ital., X, fase. II. p. 133
(sep., p. 37); and elsewhere.
2) Robertson, A., 1910, p. 230.
3) Cf. Waters, 1910, t. cit., p. 239.
4) Harmer, S. F., 1891, p. 128; cf. also A. Robertson, t. cit., p. 225.
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the sigmoid curve strongly marked, .with a branch arising from each convexity. In the first
internode shown, there are 24 zooecia, a branch originating on the right side from the fourth
zooecium of that side, and a second branch much higher from the tenth zooecium of the left
side. At about the middle of its length the fertile internode bears an ovicell, which is the
moditïed fifth zooecium of the left or inner side. The zooecium which immediately follows the
ovicell, on the same side, is displaced by the growth of the ovicell, as is usual in Crisia ; and
has a specially long peristome, which is seen, on the left margin of the internode, just distally
to the ovicell. The peristome in question is distinctly narrower than the ordinary peristomes;
a feature which I have found commonly, but not universally, in the present species. The fertile
internode has the arrangement which I regard as typical of such internodes in the present
species, and as distinguishing it from C. denticulata ; a proximal branch being developed from
the outer side of the internode, and two others, on either side, from the region of the distal
part of the ovicell.
Using the notation which I have employed in my previous paper, the part of the zoarium
shown in fig. 1 may be represented by the formula :) : —
(a) Slide 107. B.2 — (24 + r4+ io'j + (iS + r4 + S' +*)
I
(9 + Ov. -f 15 + r3 + r. + y + x)
the full number of zooecia being stated for the second main internode, which is only partially
represented in the figure.
The colony of which a part is here shown is remarkable for the facts ; — that most
of the internodes are of great length, the number of zooecia varying from 22 to 32; that most
of them have an even number of zooecia; and that they usually give rise to two branches.
The examination of other specimens shows, however, that these relations are by 110
means universal. This is brought out by the following formulae :
(b) Slide 107. R.2 (another branch) — (19 + 7') -f (21 + r.) + (19 + 7') + (19 + ;-,)
(c) Slide 107. B.2 (another branch) — (il + r5) -f (24 + qr + /-,„) + (15 + 5')
(d) Slide 313. A — (s + 2r) + (7 + rJ + (9 + 4r) + (9 + r,) + (9 + 2r) + (11 + rj
I
(11 + 2'- + r3) +
In these formulae most of the internodes have a single branch and an odd number of
zooecia; but in [c] one of them agrees with (a) in having two branches and an even number.
This seems to be the general rule, but it is not universal, as is shown by (d), where an odd-
numbered internode bears two branches.
When only a single branch occurs, its position varies, usually from the 2nd to the 7th
zooecium of its own side. The formula (d) represents a main stem, attached by rootlets at its
base. It illustrates the tendency which has already been pointed out of forming at first short
internodes with few zooecia and the branch arising as low as the 2nd zooecium. These small
1) The limits of the internodes are indicated by the brackets. The numerals denote the number of the zooecia in an internode:
Ov = ovicell: x = growing point: rt = a branch given off by the 4th right zooecium: 10' = a branch from the tenth left zooecium.
Juli.ikn and Cai.VET (1903, p. 109) consider this a useless rnethod of notation; but it appears to me a convenient method of recording
some of the characteristic features of the species.
IOO
IOI
internodes are of a form entirely different from that of the yoimger internodes; being narrow
and with the peristomes hardly developed: — which may, however, indicate that the projecting
portions have been lost with the atrophy of the corresponding polypides. But the reference to
the present species is justihed by the fact that the more peripheral internodes of the same stem
take on a broad form and may become fertile internodes bearing ovicells of the typical character.
The number of zooecia in the internode remains low in this colony, even in the peripheral parts.
The ovicell l) of the present species has the form shown in figs i, 7. lts inflated part
begins very suddenly, and the most prominent region is situated near its distal end. The
ooeciopore is of the form found in C. dentictilata, and is a small transverse slit, visible only
when the ovicell is seen from its distal end (fig. 6). In a single case I have found twin-ovicells
(fig. 8), similar to those described by d'Orbigxy ~) in C. patagonica. In rare cases, apparently
where the growth is not very vigorous, the ovicell is less inflated distally, and may be simply
pear-shaped.
The number of branches borne by the fertile internodes is typically three, as shown in
fig. i; but one of these may be suppressed; and in a single instance (fig. 4) I have found
four branches.
I have referred the specimens from Stat. 77 with some doubt to the present species.
In slide 37. R.1 the ovicells are delicate and pear-shaped, not specially inflated. This may be
due to the fact that they have been formed in parts of the erect sterns, near their origin from
rootlets, which are still slender and have not assumed the adult characters. The specimens in
question have a considerable resemblance to C. denticulata, var. gracilis, described by Busk
from the Philippine Islands 3).
The following notes may be added with regard to the specimens, recorded above, which
are not from the 'Siboga' Collection. They serve, in some cases, as a justification of the synonymy,
where references to the Memoirs cited will be found.
The specimens from Torres Straits agree in all essential respects, including the characters
of the ovicells, with the 'Siboga' material.
The same may be said of the Oueensland specimens. The material obtained from the
Australian Museum consisted of unsorted fragments which were part of the remains of the
Collection described by Prof. Haswell. There is thus a strong presumption that the Crisia
belongs to the species described by that author as C. terrac-rcginac. In his account of this
species Haswell gives the number of the zooecia in an internode as 16 — 22, the complete
internode figured having 18. The occurrence of an even number of zooecia is somewhat unusual *)
in Crisia. Haswell further states that the branches arise from the 8th to the i3th zooecium
(i. e. from the 4th to the 7th of one side); and that the joints are black. It may be noted, in
1) It is uncertain to what extent the absence of ovicells indicates that a colony is of the male sex. Miss Robertson' (1903,
"Embr. Crisia" Univ. Calif. Publ. Zool. I, p. 116) states that C. eburnca (= C. occidcntalis Trask, cf. Robertson, 1910, p. 239), is
"certainly dioecious", fhough this did not appear to be the case in all the species investigated.
2) d'Orbigny, A., 1839, "Voy. Amér. Mérid.", T. V, 4e Partie, "Zooph.", PI. I, fig. I : — see also PI. XII, fig. 13 in my
paper on Crisia, already referred to.
3) Busk, G., 1886, l.cit.
4) In C. cribraria Osburn (1912, p. 216) describes the internodes as normally constituted of an even number of zooecia.
IOI
102
this connexion, that in the original account of C. elongata, the only complete internode figured
by Milne Edwards has 20 zooecia. The specimens described by that author are believed to
have come from the Red Sea.
The specimen from Lifu recorded by Miss Philipps as C. denticulata has no ovicell, but
in other respects appears to agree with the present form.
The Japanese specimen has no ovicells, but is referred with some doubt to the present
species. C. nigrijuncta was described by Ortmann from Japan-, and, as its name indicates, it
possesses black joints.
The 'Challenger' specimen, from off Fiji, has no ovicells, but in other respects agrees
closely with the 'Siboga' material. The internodes have a pronounced sigmoid curvature, and
some of them have two branches and an even number of zooecia, in one case rising as high as
36. The joints are brown ; but it may be noted that this specimen differs from others recorded
in coming from much deeper water (1450 fathoms).
The specimens from the Red Sea determined by Mr Hincks as C. elongata are mere
frao-ments; and although the determination appears to be probable, I cannot be confident
about it.
I have not examined specimens from the Sudanese Red Sea; but there can be little
doubt that the form recorded by Mr Waters as C. denticulata is the present species. I have
found the "thin oval spot" recorded by that author in Canada balsam preparations ('Siboga',
N° 253. A.2; Queensland). It occurs, in the position shown by Waters in PI. XXV, fig. 11, on
the frontal side of the zooecium, just where the peristome turns frontally to become free. I have
not been able to make out its significance, but it is possible that it may be a result of the
sudden bending of the wall of the zooecium at this point. The "narrow tube from the distal
base of the ovicell" described by Waters (p. 233, PI. XXIV, figs 1, 2) is clearly the modified
peristome of the zooecium which next succeeds the ovicell, as described above, p. 100, and
shown in fig. 1 .
In his most recent paper (19 14) Waters records both C. denticulata and C. elongata
from the neighbourhood of Zanzibar; and both are said to have the "small mark" at the base
of the peristome. C. elongata is described as having only one branch on each internode,
springing from the 6th to the ioth zooecium of one side. I have shown above that the number
of branches is variable in the 'Siboga' material; and I think it not improbable that the two
forms recorded by Waters really belong to one species. This author states that ovicells are
not known in C. elongata.
Measuremen ts, in ,u, of 'Siboga' specimens: —
Diameter of orifices, 50 — 75 ;
From centre of orifice to centre of the next orifice, on the same side of the branch, 275 ;
Width of branch, 250 — 300 ;
Length of ovicell, measured from the distal end of the preceding zooecium, on the same side
of the branch, 500 — 750;
Width of ovicell, 500 — 575.
2. Crisia cuneata Mapl. (PI. VIII, figs 13 — 17).
Crisia cuneata Maplestone, 1905, "Lord Howe Island Pol.", Proc. R. Soc. Vict. (N. S.) XVII,
p. 390, PI. XXIX, fig. 12.
? Crisia cylindrica Ortmann, 1889 (nee Busk), "Japan. Bry.", Arch. f. Naturg. LVI, I. p. 58,
PI. IV, fig. 17.
Crisia circinata Waters, 1914, "Mar. Fauna Brit. E. Afr.", "Bry. Cycl.", Proc. Zool. Soc.,
p. 840, PI. I, figs 7 — 9.
120. B. Stat. 164. i°42'.5 S., I30°47'.5 E., 32 Metres; sand, small stones and shells.
and (Mus. Zool., Cambridge), Japan, off Tokyo, 40 fathoms, A. Ow'STON Coll., Reg. June 23,
1902, Slide 5. AM. (Now transferred to the British Museum).
Zoarium small and delicate. Margin of branches strongly serrate, owing to the marked
projection of the zooecia, the peristomes of which are directed obliquely outwards. The zooecia
of each pair are in contact in the middle line of the branch. Internodes variable in length,
sometimes consisting of 5 or 7 zooecia, but commonly more elongated. Branching rather irregular,
the branches arisinsr at variable heio'hts in the internode.
(Japanese specimens) Ovicell projecting strongly in a direction at right angles to the
branch, with a relatively long, curved ooeciostome arising not far from the point where it
becomes free.
I have found only minute fragments, without ovicells, in the 'Siboga' Collection ; and they
were unfortunately damaged after being mounted. I should not have ventured to record the
species if it had not been for the evidence of a Japanese specimen in the Cambridge Collection,
agreeing well in zooecial characters with the fragments from Stat. 1 64, except in being more
robust; and hearing ovicells of the remarkable type shown in figs 15 — 17.
C. cuneata was described by Maplestone from Lord Howe Island, E. of New South
Wales. The ovicells described and figured in the original account are of the type figured in
the present Report, but they are said to have "a fiattened distal end on which is an oval
aperture". This appears to indicate that the calcification of the ovicell was not complete; and
that the ooeciostome is not described in the original account.
The ovicells of the Japanese specimens (figs 15 — 17) are remarkably prominent, their
distal extremity being in the form of a cylinder terminated by a rounded or conical end, the
elongated being at right angles to the surface of the branch. The ooeciostome is a conspicuous
tube, of slightly curved form ; and it originates, in the usual position, not far from the point
where the ovicell becomes free. Ovicells of a similar shape have been described in other species
of Crisia and particularly in the forms referred by Busk respectively to C. edwardsiana d'Orb. l)
and C. biciliata MacGill. 2), and in C. injlata Waters 3) ; but in these cases the ooeciostome is
given off near the end of the produced part of the ovicell ; while in the first two species the
zooecia bear spines.
A portion of the zoarium of the 'Siboga' material is shown in fig. 13. It is unfortunately
too incomplete to enable any positive statements to be made with regard to the mode of
1) Busk, G., 1875, PI. II, fig. 7. See also Miss A. Robertson, 1910, p. 237, PI. XIX, fig. 10.
2) Ibid., 1S86, Challenger Rep., Pt L, PI. I, fig. I c.
3) Waters, A. W., 1914, PI. I, fig. 2.
103
104
branching. It was, however, noticed before the specimen was damaged that branches were given
off from the 2"d or 3rd zooecium of one side, or higher in the internode; and that two branches
might be given off by the same internode, on the same or opposite sides. The basis ra mi
is short, usually not reaching the zooecium on the proximal side of the one with which it is
connected. The internode is narrow, the two zooecia of a pair being in contact with one another
medianly along the whole length of their "immersed" part. The peristome is of some length,
and is directed somewhat obliquely, so that the outline of the internode appears strongly serrate.
The proximal part of the colony in the Japanese specimen (fig. 14) has so close a
resemblance to the 'Siboga' specimen that there is good reason for referring it to the same
species. The zooecia are, however, larger, and the basis rami is long enough to reach the
preceding zooecium. The specimen shown in this figure is an incomplete internode, which bears
two branches on the left side, foliowed by a third branch on the right side. An ovicell is
commencing to develop at the distal end of the principal internode. The remainder of the
Japanese specimens consist of isolated internodes, some of which are fertile. In one case three
branches arise from one side of an internode ; and some of the internodes cannot have had less than
30 zooecia. When seen in side view (fig. 1 6) the zooecia are observed to overlap one another
to a considerable extent, and in a way very different from that characteristic of C. elongata.
In the latter a zooecium does not overlap its predecessor on its basal side, but it extends for
a considerable distance on its inner side down the middle of the branch. The internode is thus
broad as well as being flattened. In C. cuneata the branch is of considerable fronto-basal depth,
and it appears narrow when seen in frontal view, in consequence of the way in which the
zooecia overlap. In some of the internodes of the Japanese material, obviously belonging to
the peripheral parts of a colony, the peristomes are much elongated.
C. cylindrica was originally described by Busk *) from Tristan da Cunha. The present
species agrees with it in its narrow branches and in the median contact of the zooecia of the
same pair when seen in frontal view. The elongated internodes and the mode of branching
seem also to be very similar in the specimens from the three localities under consideration. It
thus apjiears possible that the 'Siboga' specimen and that from Japan should be referred to
C. cylindrica. But in the absence of ovicells in Busk's fio-ures, and taking- into consideration
the distance which separates Tristan da Cunha from the localities considered in this Report, I
think it would be unsafe to assume that the specimens here described belong to Busk's species.
I have therefore referred them to C. cuneata, in which the ovicell has been described, even
though imperfectly, and which comes from a nearer locality. The Japanese form referred by
Ortmann to C. cylindrica is insufficiently described and figured ; but in view of the practical
identity of locality in the two cases, the Japanese specimens in the Cambridge Collection may
fairly be considered to belong to the same species as Ortmann's. Waters ") has recorded C.
cylindrica, though with some hesitation, from the Sudanese Red Sea; and I should have been
inclined to regard this as belonging to the same species as the Japanese form if it had not
1 i Bi >K, (1., 1886, p. 7, PI. II, figs 2, 2«, 4, 4<7.
j Waters, A. W., 1910, p. 235, PI. XXV, figs 14, 15.
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been for the account of the ovicells, which are said to be "shortly pyriform". Waters' figure
(PI. XXV, hg. 15) gives little indication of the remarkable elongation of the ovicell which is
so characteristic a feature of the Japanese specimens.
Mr Waters :) has just described, under the name of C. circinata, a species from Zanzibar
which he thinks may be identical with C. cuneata Mapl. It appears to me probable that this
form and those here recorded belong to the same species, as is shown particularly by the form
of the ovicell and of the ooeciostome.
Measurements, in u. : —
Diameter of orifices, 90 ('Siboga'); 120 (Japan);
Distance apart of zooecia of the same side of the branch, 300 ('Siboga'); 240—400 (Japan);
Width of branch, 145 — 200 ('Siboga'); 200 — 250 (Japan);
Length of ovicell (projection from frontal surface of branch), 650 — 700 (Japan);
Diameter of ovicell, 350 (Japan).
o-
Crisia kerguelensis Busk. (PI. VIII, figs 9 — 1 1).
Crisia kerguelensis Busk, 1876, "Descr. Pol. Kerguelen's Island", Ann. Mag. Nat. Hist. (4)
XVII, p. 117.
Crisia kerguelensis Busk, 1879, "Zool. Coll. Kerguelen's Land", "Pol.", Phil. Trans. CLXVIII,
p. 197, PI. X, figs 17, 18.
Crisia kerguelensis Studer, 1879, "Fauna Kerguelensland", Arc'h. f. Naturg. Jahrg. XLV, Bd I,
pp. 124, 133.
Crisia kerguelensis Studer, 1889, "Forschungsreise Gazelle", III Theil, pp. 140, 145, 158.
> Crisia eburnea, var. laxa Busk, 1886, Challenger Rep., L, p. 4, PI. II, figs 1, \a (Kerguelen).
37. S. Stat. Jj. Borneo Bank, 3°27'S., ii7°36'E., 59Metres; fine grey coral-sand (fragments).
41Ó. K. Stat. 321. 6°S'.SS., U3°30'E., 82 Metres; fine grey mud (one fragment).
Zoarium delicate, the branches narrow and the zooecia of the same pair in lateral contact
with one another on the frontal side of the internode. Lower internodes consisting of a single
zooecium, the number increasing from the proximal end of the colony and becoming fairly large
in the peripheral internodes. Internodes with an odd or even number of zooecia, a branch
commonly given off by the lowest zooecium, and another, on the opposite side of the branch,
from the second zooecium of that side. Basis rami very short, not nearly reaching the
preceding node or zooecium. Peristomes long, abruptly bent frontally at their base. Ovicell with
its inflated region entirely free and set at an acute angle to the surface of the branch ; the
ooeciostome near the distal end, on the side facing the fertile internode.
One ovicell is present (tig. 11) on the specimen from Stat. jj. It agrees with those
described by Busk sufficiently well to make the determination probable.
At the proximal end the present form has considerable resemblance to C. geniculata, a
species which C. kerguelensis was said to resemble by Busk. This is shown in fig. 9, where
the first five internodes consist each of a single zooecium. The next two internodes in the same
1) Waters, A. W., 1914; see synonymj'.
SIBOGA-EXrEDITIE XXVIII a. 1A
io6
specimen consist of three zooecia each, and the number then rises to a much larger one. The
formula of this fragment is as follows: —
(i) + (i) + (i)+(i)+(i) + (3) + (3+ k) + (ii +r, +2r + r5 + x).
Another, on the same slide, is: —
(3) + (5 + i' + r,) + (6 + r, + 2>-) + (2 + ;-, + x).
In hg. 10 it will be noticed that two of the internodes have an even number of zooecia
(4), and that the first of these two has no branch, while the second produces a branch from
its lowest zooecium. One of these internodes shows the elongated peristomes, angulated at their
base, which seem to be characteristic of the present species. The formula of the fragment
from Stat. 321 is: —
(6 + ir + r2) + (6 + v + r,) + (6 + v + ;-.,) + (1 + v + x)
(9 + 2') + (3)
(*)
(5 + 2'- + *)
(6 + ;-2) + (.v) (2H
The single ovicell (fig. 11) is of the same general type as in C. cuncata, but it is
considerably smaller. It is seen from the side, a position which shows the free character of its
inflated region to aclvantage. The ooeciostome is short but distinct, and it is produced into a
point beyond the ooeciopore.
M easuremen ts , in fj.: —
Diameter of orifices, 60 ;
Distance apart of zooecia, of the same side of the branch, 300 — 400 ;
Width of branch, 100 — 150;
Length of ovicell (projection from frontal side of branch), 335;
Width of ovicell, 170.
4. Crisia geniculata Milne Edwards. (PI. VIII, fig. 12).
Crisia geniculata Milne Edwards, 1838, "Mém. Crisies", Arm. Sci. Nat., Zool., (2) IX, p. 197,
PI. VI, figs 1— ie.
Crisia geniculata Harmer, 1891, "Brit. Spp. Crisia", Quart. J. Mier. Sci., XXXII, p. 172,
PI. XII, figs 7, 8.
Crisia geniculata Duerden, 1S93, "New and rare Irish Pol.", Proc. R. Irish Acad. (3) III,
p. 126.
Crisia geniculata Jullien & Calvet, 1903, "Rés. Camp. Prince de Monaco", XXIII, p. 1 10.
Crisia geniculata Robertson, 19 10, "Cycl. Bry. W. Coast N. America", Univ. Calif. Publ.,
Zool., VI, N° 12, p. 235, PI. XVIII, figs 6, 7; PI. XIX, fig. 8.
Filicrisia geniculata D'Orbigny, 1853, "Pal. Frang. Terr. Crét." V, p. 604.
Crisidia cornuta, var. geniculata Busk, 1S75, "Cat. Mar. Pol. Brit. Mus.", III, p. 3, PI. I,
figs 1—4.
Crisia cornuta, var. geniculata Hincks, 1SS0, "Hist. Brit. Mar. Pol.", p. 419, PI. LVI, fig. 4.
Crisia cornuta, var. geniculata Calvet, 1902, "Bry. mar. Cette", Trav. Inst. Zool. Montpellier,
(2) Mém. n, p. 71.
(Mus. Zool., Cambridge.) Torres Straits, 88, A. C. Haddon Coll., Reg. Feb. 24, 1898 (now
transferred to the British Museum).
106
107
Zoarium delicate. Internodes consisting of one, two or three zooecia, giving off a branch
on one or both sides. Peristomes very long and delicate, not much less than half the length
of the entire zooecium. Ovicells wanting.
The specimen described is from Torres Straits, and I have not found this species in the
'Siboga' Collection. In its zooecial characters it agrees closely vvith European specimens, except
that the peristomes are perhaps rather longer. In the absence of ovicells the determination
cannot be regarded as certain.
By most authors C. geniculata lias been regarded as a form of C. cornuta. I have
attempted, in an earlier paper !), to show that this view is incorrect, and that the species is
characterised by the relations of the ovicell 2), which is not free distally but is adnate on its
basal surface to one or two zooecia. As no ovicells are present in the Torres Straits specimen
it is impossible to say whether the form here considered would have been found to agree with
European specimens in the characters of the ovicell.
The zooecial characters are illustrated by fig. 12. The internodes commonly consist of a
single zooecium, though two zooecia may occur, and the internode at the upper end of the left
side of the figure has three, the largest number observed. An internode may give off one or
two lateral branches, in the latter case not quite at the same level, even in internodes consisting
of a single zooecium. The basis rami is short and does not reach the preceding joint. An
axial joint is formed in such a way as to leave the proximal part of the zooecium across which
the joint runs as a sort of basis rami which is situated in the anode between the two divereine
peristomes of the distal zooecia (where the internode consists of more than a single zooecium),
as shown in the lowest internode of the figure. The peristomes are of great length and are
curved both in a frontal and in a lateral direction.
Measurements, in u. : —
Length of zooecia, 700 — 950 ;
Width of zooecia, 50 — 60.
Fam. Extalophoridae Reuss.
Entalophoridea (pars), Reuss, 1869, "Pal. Stud. alt. Tertiarsch. Alpen"', II, Denkschr. k. Akad.
Wiss. Wien, math.-naturw. CL, XXIX, p. 2S5.
Entalophoridae Pergens & Meunier, 1887, "Faune Bry. Garumniens Faxe", Ann. Soc. Roy.
Malacol. Belg. XXI, Ann. 1886, p. 19 = 201.
Entalophoridae Gregory, 1896, "Cat. Foss. Bry. Brit. Mus.", "Jurassic. Bry.'', p. 137.
Pustuliporidae auctt.
Idtnoneidae (pars), auctt.
Entalophora Lamx.
Entalophora Lamouroux, 1821, "Exp. Méthod.", p. Si.
Entalopliora Lamouroux, 1824, "Encycl. Méthod.", "Zooph.", p. 122.
1) l. cit.
2) Miss Robertson, t. cit., p. 234, desenbes a transverse septum passing across the ovicell in its proximal half, in this and
certain other species of Crisia.
107
ioS
Entalophora Gregory, 1896, t. cit., p. 137.
Pustulopora and Pustulipora auctt.
BuSK and others have used Pustulopora or Pustulipora in place of Entalophora. It has,
however, been pointed out by Gregory (1. cit.) that Pustulopora, as originally introduced by
de Blainville l) was a synonym of Spiropora Lamouroux, 182].
The genus Entalophora includes a number of recent and fossil species in which the
zoarium is erect and the peristomes open irregularly on all sides of the sterns. A key to the
Cretaceous forms has been published by Lang3). Gregory (t. c, p. 217) considers that the
Entalophoridae are nearly allied to the Diastoporidae, partly on the evidence of a specimen
of Entalophora in which the erect stem grows from a Beremcea-like encrusting portion. Some
of the names have been used for both recent and fossil species ; but it is extremely clifhcult
to decide how far this procedure is justifiable.
1. Entalophora proboscidca M. Edw. (PI. X, fig. 12).
Pustulopora proboscidea H. Milne Edwards, 1838, "Mém. Crisies", Ann. Sci. Nat. (2) IX,
p. 219, PI. XII, fig. 2 (recent, Mediterranean).
Pustulopora proboscidea Busk, 1875, "Cat. Mar. Pol. Brit. Mus.", III, p. 21.
Pustulopora proboscidea Haswell, 1879, "Cycl. Pol. Port Jackson", Proc. Linn. Soc. N. S. Wales,
IV, p. 352.
Pustulopora proboscidea Busk, 1886, Challenger Rep. ;1), L, p. 19, PI. IV, figs 1 — ib. (Heard
Island, Prince Edward Island, S. Indian Ocean).
Pustulipora proboscidea Haswell, 1880, "Pol. Queensland", Proc. Linn. Soc. N. S. Wales, V,
p. 35 (110 description).
Entalophora proboscidea Gregory, 1899, "Cat. Foss. Bry. Brit. Mus.", "Cret. Bry.", I, p. 221
(Naples, recent).
Entalophora proboscidca Waters, 1904, "Rés. Voy. Belgica", "Bry.", pp. 9, 91, PI. IX, figs \a,
\b (Antarctic and Cosmopolitan).
Entalophora proboscidea Norman, 1909, "Pol. Madeira", J. Linn. Soc. Zool., XXX, p. 2S0,
PI. XXXV, figs 1—3.
Entalophora proboscidea Thornely, 1912, "Mar. Pol. Ind. Ocean", Trans. Linn. Soc. (2) Zool.,
XV, p. 156 (Amirante and Seychelles Islands; 110 description).
': Entalophora indica D'Orbigny, 1853, "Pal. Frang. Terr. Crét.", V, p. 781 (Straits of Malacca).
This list of synonyms might be greatly extended, but most of the references would be
very doubtful. A number are given by Miss Jellv *). Gregory, t. cit., p. 219, includes many
of the citations of E. proboscidca under E. virgula, a Cretaceous species which he regards as
nearly related to the recent E. proboscidca (pp. 217, 221).
329. A. Stat. 64. Kambaragi Bay, Tanah Djampeah, o — 32 Metres; coral, coral-sand.
239. A. Stat. 310. 8° 30' S., U9°7'.5E., 73 Metres; sand with a few pieces of dead coral.
(? sp.) 449. B. Stat. 59. W. entrance Samau Strait, 390 Metres; coarse coral-sand with small stones.
251. H.1 Stat. 315. Anchorage E. of Sailus Besar, Paternoster Islands, o — 36 Metres; coral
and Lithothamnion.
ij de Blainville, II. M. I).. 1S30, Dict. Sci. Nat. LX, p.382; and 1834, "Man. d'Actinol.", p. 418 {Pustulipora in Index, p. 691).
2) Lang, W. D., 1906, "Key publ. Figures Cret. Entalophora", Geol. Mag. (N. S.), Dec. V, III, p. 462.
3) In the synonymy liere given BUSK statcs that the [right hand] figure on "PI. XVIII. A" (errorim, for XVII. A.) of the
British Museum 1 atalogue does not belong to MlLNE EDWARDS' species, and implies that il was taken from a Shetland specimen.
4) J 1:1.1. v. E. C, i8Sg, "Syn. Cat. Ree. Mar. Bry.", p. 89 (under /•:. raripord).
108
109
Also (Mus. Zool., Cambridge) : —
Torres Straits, 195, A.C. HADDON Collection, Reg. Feb. 24, 1898.
Queensland, Port Denison, Miss E. C. JELLY, Reg. May 24, 1895.
Oueensland, Port Denison and Holborn Island, Australian Museum, Sydney, Reg. Oct. 23,
1S99 (from remains of the Collection described by Prof. W. A. Haswell).
Zoarium consisting of a few coarse, irregularly arranged branches which bifurcate rather
seldom and do not keep in one plane. Zöoecia few and large, often with greatly produced
peristomes. Ovicell occurring at a bifurcation, variable in form, but extending some way up
one of the branches thus formed. Ooeciostome terminal, meeting the rest of the ovicell at an
angle, and when fully developed with a transversely elongated ooeciopore, bounded by well
developed lips.
The present species is distinguished from the other forms dredged by the 'Siboga' by
its more robust zooecia, and by the characters of its ovicell. lts growth is very irregular, the
branches not keeping in the same plane, and varying in thickness. The largest colony examined
(239. A.) is about 20 mm. in greatest length, and consists of two main lobes, which diverge
from one another close to the base of the colony at an angle of nearly 1800. Not more than
three or four bifurcations are present in any branch, so that the colony does not attain much
complexity of growth.
The zooecia, while maintaining a practical uniformity in their sectional area, vary much
in other respects. The peristome is either short or long; and it may continue the direction of
the proximal part of the zooecium in a uniform curve, or it may leave that part at a distinct
angle. The direction of its growth may be uniform or irregular, an alteration of the curve
being frequently visible at some point in its length. It is frequently marked by circular Unes of
orowth. As will be seen from fig. 12, the number of zooecia present in any transverse section
of a branch is small compared with that found in some other species. This character depends,
ho wever, on the degree of vigour in the growth of the branches, some of which may be
slenderer than in the portion figured; while in other parts, particularly in the region immediately
preceding a bifurcation, the number of zooecia present at the same level may be increased.
The ovicell of the specimen figured (fig. 1 2) commences at a bifurcation and extends
along only one of the branches so formed. This fertile branch shortly bifurcates again, although
one of the branches has been broken off. With the first bifurcation placed in the plane of the
drawing the ovicell is seen from the side; but if the bifurcation which occurs at about the
middle of the ovicell had been in that plane it is obvious that the ovicell would have been
seen in frontal view. This is a consideration which has to be borne in mind in comparing this
specimens with others. At the point where the second bifurcation was formed there has been
an interruption in the continuity of the growth of the ovicell, as is shown by the persistence
of part of the margin of the growing end of the branch as the ridge which is seen crossing
the ovicell. The remainder of the roof of the ovicell has been formeel by a calcareous lamina
developed slightly within this edge. Another sudden change in the direction of growth is
indicated by a kind of collar present near the base of the ooeciostome.
The ooeciostome is a wide tube which leaves the ovicell at its distal end; and it is
109
I IO
directed somewhat frontally, with a distinct angle at its junction with the ovicell. The ooeciopore
is transversely elongated and it is bounded by fairly distinct lips.
The ovicells of the specimens referred to the present species differ from one another;
luit this appears to depend largely on the nature of the growth of the fertÜe branch. In another
specimen (239. A.1) from the same Station the ovicell is developed from a short thick branch,
which shows only a slight indication of bifurcating in the region of the ovicell, which is thus
considerably wider at its distal end than in the specimen shown in fig. 1 2 (the ovicell having
shared in the general widening of the branch which has taken place in this region). The
ooeciostome has, however, the same terminal position, although it is not quite so well developed
as in the branch figured.
In the specimens from Queensland three ovicells are present, all of a more elongated
type than in fig. 12. One of these begins at a bifurcation and extends for some distance up
the inner side of one of the branches so formed. The branch in question is a slender one, and
the ovicell is correspondingly narrow. The other two ovicells begin before the bifurcation and
extend along one of the branches. In one of them the ovicell is on the side of the branch
facing the other branch, and is accordingly seen in side view with the bifurcation in the plane
of the slide. The other ovicell is quite similar, except that it is seen in frontal view with the
bifurcation in the same position. In all these cases the ooeciostome is terminal and is set at an
angle to the rest of the ovicell.
The other specimens recorded above have no ovicells ; but, from their zooecial characters,
329. A. and the specimen from Torres Straits may be placed with the present species with
some confidence-, while 251.H.1 is probably correctly determined and 449. B. — a fragment in
poor condition — is doubtful. Two specimens dredged by the 'Challenger' off the Marion Islands,
in the Cambridge Collection, and referred to E. proboscidioides, have some resemblance to the
'Siboga' specimens ; but their sterns are more regularly cylindrical ; and in the absence of ovicells
it would be unsafe to assume that they belong to the present species.
Measurements, in <j. :
Width of branch, at lower end of fig. 12, 850 ;
Length of longest peristome, fig. 12, 1,400; diameter, 220 — 250;
Length of ovicell, fig. 12, 2,280;
Width of ovicell, at proximal end, fig. 12, 780;
Width of ooeciostome, fig. 12, 250 ;
Diameter of orifices, fig. 12, 150 — 200.
Gregory (t. cit., p. 222) gives the diameter of the orifices of a recent specimen from
Naples as 210 u..
2. Entalophora delicatula Busk. (PI. X, fig. 11).
Pustulopora delicatula Busk, 1S75, -'Cat. Mar. Pol. Brit. Mus.", III, p. 20, Tl. VLB, fig. 3
(Queensland).
? Pustulipora delicatula Haswell, 1SS0, "Pol. Queensland", 1'roc. Linn. Soc. N. S. Wales, V, p. 35.
1 10
I I I
? Entalophora delicatula Ortmann, 1S89, fcJapan. Bryozoenfauna", Arch. f. Naturg., Jahrg. LVI,
Bd I, p. 61, PI. IV, figs 28 a, 2%b.
Entalophora delicatula Philipps, 1899, WlLLEY's "Zool. Res.", Pt IV, pp. 441, 449 (Lifu).
> Pustulipora fragilis Haswell, 1880, t. cit., p. 35, PI. I, fig. 2.
? Entalophora fragilis Waters, 1SS7, "Bry. X. S. Wales", III. Arm. Mag. Nat. Hist. (5) XX.
p. 259.
Entalophora deflexa Smitt (nee auctt.), 1872, "Floridan Bry.", I, K. Svensk. Handl., X,
N° 11, p. 11, PI. V, figs 28—30 (fide WATERS).
Entalophora wasinensis Waters, 1914, "Mar. Bry. B. E. Afr.", "Bry. Cycl.", Proc. Zool. Soc,
p. 840, PI. II, figs 1 — 4, 9; text-fig., p. S41 (Zanzibar, etc., 10 fathoms; Florida, SMITT).
376. L. Stat. 213. Saleyer Anchorage and surroundings, o — 36 Metres ; coral-reefs, mud and
mud with sand.
298. B. Stat. 273. Anchorage off Pulu Jedan, E. coast of Aru Islands, 13 Metres; sand and shells
(on 298. A., Amathia convoluta Lamx).
217. A. Stat. 301. Pepela Bay, E. coast of Rotti Island, io°38'S., I23°25'.2E., 22 Metres;
mud, coral and Lithothamnion.
(sp.incert.) 57. X. Stat. 80. Borneo Bank, 2°25'S., U7°43'E., 40—50 Metres; fine coral-sand.
The following specimens, in the Collection of the University Museum of Zoology, Cambridge,
are referred, somewhat doubtfully, to the same species :
Japan, from rooting spicules of Euplectella marshalli, Prof. K. MlTSUKURI, Reg. Sep. 23, 1S96.
Queensland, Port Denison (two slides), Miss E. C. JELLV, Reg. May 24, 1S95.
Queensland, Port Denison, from remains of the Collection described by Prof. W. A. HASWELL,
Australian Museum, Sydney, Reg. Oct. 23, 1899.
Lifu, Loyalty Islands, A. WlLLEY Collection, Reg. Mar. 1, 1898 (determined by Miss Philipps
as E. delicatula Busk.
Torres Straits, 132, 190. A. C. HADDON Coll., Reg. Feb. 24, 1898.
Zoarium consisting of delicate branches. Zooecia small, the peristomes long and prominent;
usually about four series visible in one half of the branch. Ovicell more or less pyriform ; if
present at a bifurcation extending into one of the branches only : its roof not traversed by
zooecia. Ooeciostome terminal, with a transversely elongated ooeciopore, bounded by lips which
are variable in their development.
If the specimens of which a list is given above are correctly referred to one species,
there is a considerable amount of variation in the characters. The branches and the zooecia
are much slenderer than those of E. proboscidea, as is seen by comparing fig. 1 1 with fig. 1 2
(both drawn to the same scale). In one or two cases, particularly in the specimens from
Queensland, some of the branches are very straight; and, except for the projection of the
peristomes, cylindrical in form '). In others, as in 376. L., these features are not so pronounced.
The peristomes are usually long and slender. The ovicell differs from that of the species next
to be described in not having its roof traversed by zooecia ; although some of the peristomes
at its sides may be laterally adnate to its roof.
The ovicell, although variable in shape and in its position with relation to a bifurcation,
appears to have some features which are fairly constant. If developed in the region of a
bifurcation, it does not divide with the branch, but extends into only one of the branches (fig. 1 1).
1) These specimens appear to resemble Pustulipora fragilis Haswell (1. cit.).
II I
I 12
As is usual in Cyclostomes, its pores are more numerous than those of the zooecia; and they
are also distinctly larger. The ooeciostome is terminal. In the specimen figrired it is seen
sideways; although, as a second bifurcation is formed at its level, it would have been seen
in frontal view [f the branch had been arranged with the plane of the second bifurcation
horizontal. In other specimens in which no second bifurcation of the fertile branch takes place,
the ooeciostome is situated symmetrically with regard to the proximal end of the ovicell. which
is more or less pyriform. In fig. 1 1 the distal lip of the ooeciostome is seen to be curved over
the ooeciopore, as a sort of hood, thus resembling the ooeciostome of Tiibulipora phalangea ;
and its proximal lip is hardly developed. In some of the other specimens the distal lip does
not have this hood-like character; and the ooeciopore is transversely oval, with its proximal
and distal lips about equally developed. The type-specimens (Brit. Mus. 75. 5. 29. 30) of Pustu-
lopora delicatula Busk show ovicells of the general character of those here describecl in two
of the branches.
Measurements, in u.:
Width of branch at lower end of fig. 11, 250 ; of left branch, fig. 11, 400;
Length of part of ovicell visible in the same figure, measured in a straight line, 1,950;
Greatest width of ovicell, as seen in the same figure, 450 ;
Diameter of zooecia, fig. 11, 75 — 100; of orifices, So.
The specimens here describecl appear to agree closely with E. wasinensis, which Mr Waters
regards as including E. deflexa Smitt, 1872 (nee Couch). The agreement is manifested in the
form of the zoarium, in the measurements of the various parts, and particularly in the characters
of the ovicell and of the ooeciostome. Waters particularly comments on the fact that none of
the zooecia pass through the ovicell.
3. Entalophora intricaria Busk (? sp.). (PI. X, figs 13, 14).
Pustulopora intricaria Busk, 1S75, "Cat. Mar. Pol. Brit. Mus.", III, p. 22, PI. X, figs 1, 4
(South -Australia).
Pustulopora intricaria Haswell, 1S79, "Cycl. Pol. Port Jackson", Proc. Linn. Soc. N. S. Wales,
IV, p. 352.
Entalophora intricaria Waters, 1914, "Mar. Fauna Brit. E. Afr.", ''Bry. Cycl.", Proc. Zool.
Soc, p. S42.
[ Stat. 164. i°42'.5 S., I30°47'.5E., 32 Metres; sand, small stones and shells.
I35-F.
506. B.
202. D. Stat. 282. Anchorage between Nusa Besi and the N. E. point of Timor, 27 — 54Metres;
sand, coral and Lithothamnion.
(? sp.) 106. A. Stat. 144. Anchorage N. of Salomakiëe (Damar) Island, 45 Metres; coral-bottom and
Lithothamnion.
(? sp.) 351.A. Stat. 240. Banda Anchorage, 9 — 45 Metres; black sand, coral, Lithothamnion-bank
in 18 — 36 Metres.
(?sp.) 254. F. Stat. 315. Anchorage E. of Sailus Besar, Paternoster Islands, o — 36 Metres; coral and
Lithothamnion.
The following specimens, in the Collection of the University Museum of Zoology, Cambridge,
may belong to the present species : —
Torres Straits, 45, 90, A. C. HADDON Coll., Reg. Febr. 24, 1898.
H3
Zoarium rather more robust than that of E. dcticatitla. and the zooecia slightly larger;
their immersed portions sometimes very long. Ovicell not bifurcating with a branch ; its roof
pierced by isolated peristomes, and its ooeciostome not terminal.
The present species is slightly coarser than E. delicatula, and its zooecia are somewhat
larger. In some of the specimens referred to it, the zooecia have a very long portion immersed
in the general substance of the branch, before they give off their free peristomes J) The feature
which specially distinguishes this form is the ovicell, which has the character shown in fig. 14.
In this specimen the ovicell is rather broad, with a gently convex frontal surface, which descends
very steeply into the lateral walls of the branch. It is "simple" in character, not bifurcating
with the branch. In the fact that its roof is traversed by zooecial peristomes (usually isolated)
it differs from the other 'Siboga' specimens. The ooeciostome (if this structure has been correctly
identified) is situated at a long distance from the distal end of the ovicell, instead of being
terminal, as in the other two species. In fig. 14 it seems to be the opening which is adnate
to an ordinary peristome, not far from the proximal end of the ovicell; and it is not unlike
an ordinary orifice in shape. Ovicells of a similar type are present in 202. D., in which the
ooeciostomes also have the character shown in the figured branch. One of the Torres Straits
specimens (90) has ovicells in which the roof is traversed by zooecia, although the ooeciostomes
cannot be seen with certainty. The remaining specimens in the above list have no ovicells;
and the determinations must be regarded as doubtful.
All the specimens described are small fragments.
Measureraents, in rj. (fig. 14):
YVidth of branch, at lower end, 580 ;
Length of inflated part of ovicell, 1,820;
Greatest width of inflated part of ovicell, 1,000 ;
Diameter of peristomes, 130 — 150;
Diameter of orifices, 100.
Mr Waters (1. cit.) has stated that in E. intricaria the zooecia pass through the ovicell.
This is the principal justification for the determination of the 'Siboga' specimens, which are,
however, more delicate than those described by Busk and may belong to a distinct species.
Their fragmentary condition does not permit any statement to be made as to the character of
the zoarium; which, according to Busk's original description, should form "dense intricate masses
of considerable size".
Fam. Diastoporidae Busk.
Diastoporidae Busk, 1S59, "Monogr. Foss. Pol. Crag'", Palaeont. Soa, pp. 91, 113.
Diastoporidae Smitt, 1866, "Krit. fort. Skand. Hafs-Bry.", II, Öfv. K. Vet.-Ak. Förh. XXIII,
P- 395-
I use the Family Diastoporidae in the sense in which it was understood by Busk and
1) The greatly elongated zooecia seen in some of the specimens (fig. 13) have not been found in fertile branches: and it is
possible, though not very probable, that they may not belong to the same species as the fertile branches.
SIEOGA-EXPEDITIE XXVIII17.
ii4
Smitt, for adnate or foliaceous forms in which the zooecia are to a large extent immersed,
formino- a continuous sheet, and in which the peristomes are relatively little developed. The
ovicells differ in the various species, sometimes being but little expanded, and not nnlike those
of Crisia, and sometimes being considerably broadened in a transverse direction, so as to form
conspicuous inrlations on the surface of the colony. It appears to be characteristic of this Family
that zooecia with functional polypides are restricted to the periphery of the colony, the more
centrally placed zooecia becoming occluded by a calcareous covering, the form of which may
serve to distinguish the species. By many authors the Diastoporidae are regarded as falling
within the Tubuliporidae.
Berenicea Lamx.
Berenicea Lamouroux, 1821, "Exp. Méthod.", p. 80.
Berenicea Lamouroux, 1824, "Encycl. Méthod.", "Zooph.", p. 140.
Berenicea Gray, 1848, "List Brit. An. Brit. Mus.", I, p. 142.
Berenicea Reuss, 1867, "Bry. Braunen Jura Balin", Denkschr. K. Ak. Wiss. W'ien, math.-
naturw. Cl. XXVII, I Abth., p. 4.
Berenicea Novak, 1877, "Bry. Böhm. Kreideform.", Ibid., XXXVII, II Abth., p. 96.
Berenicea Gregory, 1896, "Cat. Foss. Bry. Brit. Mus.'', "Jurassic Bry.", p. 76.
Diastopora Waters, 1914, "Mar. Fauna Brit. E. Africa", "Bry. Cycl.", Proc. Zool. Soc, p- 835.
Diastopora auctt. (nee Lamouroux, 1821, p. 42).
By most writers on recent Polyzoa the genus Berenicea has been suppressed in favour
of Diastopora. Lamouroux used the former for encrusting species, and the latter for foliaceous
forms. The history of the genera was explained by Reuss (1867), who pointed out that the
distinction between them was somewhat arbitrary. Gregory (1896) argues in favour of retaining
the distinction made by Lamouroux, on the ground of convenience. Canu l) considered that
Berenicea was inseparable from Diastopora, placing the species described by him in Diastopora.
In a later paper-) he uses Berenicea in preference to the other generic name. Normax3) maintained
that Berenicea prominens Lamx was really identical with the Cheilostome Chorizopora brong-
niartii Aud. ; although in a later paper 4) he stated that B. prominens was mentioned by him
in error, and that B. annulata was the species he intended to refer to. I am in agreement
with Waters 5) in thinking that Lamouroux' Berenicea probably referred to the Cyclostomatous
species which have been placed in it by other authors, and not to Chorizopora.
1. Berenicea sarniensis Norman. (PI. XI, figs 4, 5).
Diastopora sarniensis Norman, 1S64, "Undescr. Brit. Hydr. Act. Pol.", Ann. Mag. Nat. Hist.
(3) XIII, p. 89, PI. XI, figs 4—6 (Guernsey & Jersey).
Diastopora sarniensis Busk, 1875, "Cat. Mar. Pol. Brit. Mus.", III, PI. XXXIV, fig. 5.
Diastopora sarniensis Hincks, 1880, "Hist. Brit. Mar. Pol.", p. 463, text-fig. on p. 461,
PI. LXVI, figs 7—9.
1) Canu, F., 189S, "Ét. OviceUes Bry. Bathonien", Buil. Soc. Géol. France, (3) XXVI, p. 265.
2) Ibid., 1913, "Conti-. Ét. Bry. Foss.", Ibid., (4) XIII, pp. 268—.
3) NORMAN, A. M., 1903, "Notes Nat. Hist. E. Finmark", Ann. Mag. Nat. Hist. (7) XI, p. 569.
(i [bid., 1909, "Pol. Madeira", J. I.inn. Soc, Zool., XXX, p. 299, note.
5j Waters, A. W., 1904, "Bry. Fr.-Josef Land", II, Ibid., XXIX, p. 172.
114
H5
Diaslopora sarniensis Hincks, 1884, "Pol. Oueen Charlotte Islands", Ann. Mag. Nat. Ilist.
(5) XIII, p. 206.
Diastopora sarniensis Hincks, 1887, "Pol. Adriatic", Ibid. (5) XIX, p. 30S.
Diastopora sarniensis MacGillivray, 1887, McCov's "Prodr. Zool. Vict.", Dec. XV, p. 181,
PI. CXLVII, figs 4—4/;.
Diastopora sarniensis Kirkpatrick, 1890, "Hydr. Pol. China Sea", Ann. Mag. Nat. Hist. (6) V,
pp. 17, 22.
Diastopora sarniensis Calvet, 1902, "Bry. Mar. Corse", Trav. Inst. Zool. Montpellier, (2) Mém. 12,
p. 42.
Diastopora sarniensis Jullien and Calvet, 1903, Rés. Camp. Prince de Monaco, XXIII, p. 117.
Diastopora sarniensis Calvet, 1907, "Bryozoaires", Exp. Sci. Travailleur et Talisman, VIII,
p. 465.
Mr Waters (1887, "Tert. Cycl. Bry. New Zealand", O. J. Geol. Soc, XLIII, p. 342) has
recorded Diastopora sarniensis, var. perangusta from New Zealand, both recent and fossil; but he
states that the zooecia are only about half the size of those of typical D. sarniensis from Guernsey.
81. F. Stat. 105. 6°S'N., 1210 19' E., 275 Metres ; coral-bottom.
566. F. Stat. 204. Between Islands of Wowoni and Buton, N. entrance of Buton Strait,
75 — 94 Metres; sand with dead shells.
551. C. Stat. 310. 8° 30' S., ii9°7'.5E., fi Metres; sand with a few pieces of dead coral.
And (Mus. Zool., Cambridge) : —
Japan, Uraga Channel, off Tokyo, 80 fathoms; A. OwSTON Coll., 28. H., Reg. June 23, 1902.
Zoarium somewhat irregular in form, sometimes flabellate. Marginal lamina variable.
Zooecia quincuncial, not in radiating series, peristomes free. Occlusion of the orifices partial, in
the form of a transverse calcareous plate from which rises a short, narrow tube which remains
open at its free end. Ovicells not forming a continuous ring ; not broader than long, or not
much broader than long. Ooeciostome centrally placed, near the distal end, but looking
somewhat proximally.
The present species, which I am unable to distinguish from the European B. samiejisis,
originally described by Norman from Guernsey and Jersey, is represented by only a few specimens
in the 'Siboga' Collection. In its young state (566. F.) the colony may commence with a narrow
Sfo/nafopora-\ike portion, which after a time bifurcates, at rig'ht angles, into two lobes, at an
angle of 1 8o° to one another. In the specimen described, one of these lobes has hardly developed,
while the other remains quite flabellate (fig. 5). In a later stage of growth (fig. 4) the colony is
more discoidal, but with an irregular outline. The zooecia show none of the radial arrangement
which characterises B. lineata, but are arranged quincuncially. Their proximal parts are depressed,
as in the other species; but their distal parts rise into free peristomes, which do not become
connate. In the 'Siboga' specimens, very few of the orifices are occluded, and the marginal lamina
remains narrow. In the specimen from Japan, all the centrally placed zooecia are occluded ;
and, in correlation with this fact, the marginal lamina is broad, no doubt indicating a rapid
production of new zooecia to take the place of those which are being occluded.
The "closure" of the zooecia takes the form which is well known in British specimens;
a calcareous plate being formed across the orifice, and giving oft" a short, open tube, of small
diameter. Two such "closures" are seen in fig. 5. The small tube is generally given off from
the distal half of the calcareous plate.
n6
The ovicells (fig. 4) are not so broad as those of B. lineata \ and they show no disposition
to form a continuous ring. They are very clistinctly outlined, as their inflated part commences
suddenly, even on the proximal side. In one of the colonies on slide 551.C, the roof of the
ovicell has been broken ; permitting the observation to be made that the ovicell has a round
hole in its proximal wall, of the same diameter as a zooecium. This no doubt indicates the
regfion where the ovi^erous zooecium dilated suddenlv into an ovicell. The floor of this ovicell
is complete, the subjacent zooecia being covered by a calcareous film which conceals their
outlines. The ooeciostome is placed at the middle of the ovicell, near its distal border. It is
not terminal, as it is in B. suborbicularis\ and it slopes a little proximally (see left ovicell in
fig. 4). The ooeciostome is a short tube which is terminated by a nearly circular ooeciopore.
Several ovicells may be present in the same colony.
The Japanese specimen agrees in all respects with those from the 'Siboga' Collection,
except for the difference, which has already been pointed out, of having all its central zooecia
occluded; but the form of the "closure" is as in the other specimens.
Measurements, in u. :
Greatest diameter of colony, fig. 4, 3,400;
YVidth of ovicells, fig. 4, 500, 525, 700 ;
Diameter of ooeciopore, 50 ;
Diameter of orifices, 70.
2. Berenicca lineata 'MacGill. (PI. XI, figs 6, 7).
Diastopora lineata MacGillivray, 1S85, "Descr. New Pol.", VII, Trans. Proc. R. Soc. Vict.,
XXI, p. 96, PI. III, fig. 1.
Liripora lineata MacGillivray, 1887, Ibid. XII, Trans. Proc. R. Soc. Vict., XXIII, p. 182.
Discotubigcrai lineata Waters, 1887, "Bry. N. S. Wales", II, III, Ann. Mag. Nat. Hist. (5)
XX, p. 260, PI. VI, fig. 24.
Discotubigera': lineata Waters, 1S89, "Ovicells Lichenoporae", J. Linn. Soc, Zool., XX, p. 284,
PI. XV, fig. 5 (ovicell).
i Diastopora bicolor MacGillivray, 1885, "Descr. New Pol.", VIII, Trans. Proc. R. Soc. Vict.
XXI, p. 117, PI. V, fig. 2.
? Diastopora bicolor MacGillivray, 18S7, McCov's "Prodr. Zool. Vict.", Dec. XV, p. 180,
PI. CXLVII, figs 3, ia.
? Liripora bicolor MacGillivray, 1905, "Monogr. Tert. Pol. Vict.", Trans. R. Soc. Vict., IV,
p. 131, PI. XX, fig. 8; Hall, Ibid., Appendix, p. 138.
? Diastopora prominens Ortmann, 1889 (nee Lamouroux, 1821), "Japan. Bry.", Arch. f. Naturg.
Jahrg. LVI, Bd I, p. 64, PI. IV, fig. 38.
104. C. Stat. 139. o°ii'S., i2/°25 E., 397 Metres; mud, stones and coral.
424. A. Stat. 240. Banda Anchorage, 9 — 45 Metres; black sand, coral; Lithothamnion-bank
in 18 — 36 Metres.
Also (Mus. Zool., Cambridge) : —
Singapore, New Harbour, 6 fathoms, Dr. R. HANIïSCH, Reg. Apr. 10, 1900.
Japan, Uraga Channel, off Tokyo, 30 fathoms, A. Owston Coll., 7. X., Reg. June 23, 1902.
Each of the above localities is represented by a single specimen.
116
ii7
Zoarium circular, with a strongly developed marginal lamina, which may become free
and cup-like. Zooecia with a pronounced radial arrangement, those of the central region with
a depressed porous region passing into the projecting peristomial part. The peristomes are
connate, in radial series; at first uniserial, but more peripherally becoming biserial or triserial.
In fertile colonies the series of zooecia appear interrupted by the ovicells, but may bifurcate in
that region, so that the number of series is greater on the peripheral side of the ovicells than
in the central part of the colony. Outside the ovicells, the zooecia are usually in connate groups,
which are two or three zooecia wide. Ovicells transversely elongated ; when several are present,
forming a ring which extends round almost the entire circumference of the colony. The more
centrally placed zooecia are closed by a terminal or slightly depressed calcareous plate.
The genus Liripora was founded by MacGili.ivrav (1887) for species which differ from
Diastopora \Berenicea\ in having the zooecia arranged in uniserial or multiserial radiating rows.
It appears to me possible that L. bicolor of the same author is identical with L. lineata.
The bathymetrical range of this species is considerable (6 fathoms = 1 1 Metres, to
397 Metres), if the specimens here described are rightly referred to the same species. The
largest colony is shown in fig. 7 (424. A.), and measures 4.4 mm. in diameter, a size which
would have been slightly exceeded if the marginal lamina had been complete. The connate,
radial arrangement of the zooecia appears to be very characteristic of this species ; and this is
a sufficiënt reason for distinguishing it from B. patina Lamk, which it resembles in several
respects. The zooecia in these series are at first arranged in a single line, but become biserial
even on the central side of the ovicell, while more peripherally they are in two or three series.
The depressed parts, between the series, in the central region of the colony, are porous, and at
first sight might be mistaken for parts of the ovicells. More careful examination shows, however,
that these parts are divided by septal lines; and the specimen 104. C. (fig. 6) shows very distinctly
that these are really the edges of the walls between the zooecia, which are accordingly depressed
in the proximal part, becoming raised in their peristomial region into the constituents of the
radiating ridges. Consecutive zooecia of a series may have their depressed part on the same side
or on opposite sides of the peristomes; in the latter case an alternate disposition being indicated.
It appears to be characteristic of the present genus that the centrally placed zooecia
cease to be functional at a relatively early stage. This is indicated by the fact that they are
terminated by the well known calcareous "closure". The shape of this closing plate offers good
characters for the discrimination of the species. In the present case the "closure" is of the type
characteristic of B. patina ; consisting of a nearly terminal plate provided with a few pores,
but not raised into a tubular prolongation. It may safely be assumed that the formation of the
terminal plate is a consequence of the degeneration of the polypide; and that the only zooecia
which possess functional polypides are those which have no "closure" and are situated in the
marginal region of the zoarium x). The great development of the marginal lamina which is so
common in this genus is probably correlated with the fact that there is here an active growth
of new zooecia to replace the more centrally placed occluded zooecia.
1) I have evidence, derived from a British specimen of B. samiensis, that the central zooecia contain no functional polypides.
"7
i iS
The inflated part of the ovicell (which has been figured by Waters, 1889) commences
verv suddenly on its proximal side, and it is hardly possible to distinguish which zooecium has
given rise to it. There is, however, every probability that, as in other Cyclostomes, the ovicell
is at first a zooecium, the distal end of which becomes greatly enlarged and dilated as the
result of the development of an egg into an embryo which gives rise to numerous larvae by
a process of embryonic fission. In fig. 6 only a single ovicell is present; and it is thus easy
to distinguish its lateral outlines. The ooeciostome is in this case a short tube which rises
symmetrically from the middle of the distal border of the ovicell, terminating in a nearly circular
ooeciopore which looks frontally.
In fig. 7 the ovicells form a ring which is quite complete except at one side, where
the growth of the colony has no doubt been checked by some object on which it was growing.
Four ooeciostomes can be distinguished, each of which lies near the proximal end of a pluriserial
group of connate zooecia, representing the peripheral end of one of the series of zooecia which
started near the centre of the colony. As these peripheral groups (about 26) are more numerous
than the radial ridges (about 1 6) in the more central part of the colony, it may be concluded
that some of the series have bifurcated, but that the point of bifurcation is concealed by the
ovicell. The number of ooeciostomes present indicates that there are probably four ovicells in
specimen 424. A.; but the lateral union of the ovicells is so intimate that their limits cannot
be made out. It is not impossible that they have fused with one another laterally, so that the
cavities have become confluent; but it cannot be ascertained whether this is the case or not
without destroying the specimen.
424. A. was growing on a small Gasteropod shell, and the Japanese specimen was also
on a shell. 104. C. is on the basal surface of a Retepore (104. B.), two or three of the fenestrae
of which are obliterated by its marginal lamina.
M e a s u r e m e n t s , in u : —
Greatest diameter of colony, fig. 7, 4,400;
Greatest diameter of ovicell-ring, fig. 7, 3,300;
Radial diameter of the same ovicells, 600 — 700 ;
Width of ovicell, fig. 6, 850 ;
Diameter of ooeciopore, 80 ;
Diameter of orifices, 100.
Tubuliporidae Johnston.
Tubuliporidae (pars), Johnston, 1838, "Hist. Brit. Zooph.", Ed. 1, p. 267.
Tubuliporidae (pars), Johnston, 1847, Ibid., Ed. 2, p. 265.
Tubuliporidae (pars), Hincks, 1880, "Hist. Brit. Mar. Pol.", p. 424.
Tubuliporidae and Idmoneidae auctt.
The Family Tubuliporidae, as used by Johnston in 1838, included the two genera
Tubulipora and Di$copora\ the former comprising species of Berenicea (Diastofiora) as well as
118
ii9
species which have retained their place in Tubulipora. Discopora was represented by D. hispida
only, a species which has been placed in Lichenopora by the majority of recent authors.
In the present Report the Family Tubuliporidae is understood to include the genera
there placed by Hincks, with the exception of Diastopora and Entalophora ; and to consist,
therefore, of Cyclostomata in which the zooecia are restricted to one surface of the colony and
are commonly arranged in connate alternating series. Cancelli are absent in the majority of the
species. The ovicell is a modified zooecium which is usually much dilated in the region where
the embryos undergo their development.
The study of recent Cyclostomata is greatly complicated by the fact that this Suborder
is represented by an immense number of fossil forms, which have frequently been named from
imperfect fragments-, and by the fact that it has been investigated to a large extent independently
by Palaeontologists and students of the recent species. The conclusions of these two sets of
workers referring to nomenclature have often been very different; and there have been many
divergences of opinion with regard to the propriety of identifying recent with fossil species. It
must be remembered that the recent species constitute but a small fraction of the number of
those known in a fossil condition. A satisfactory decision as to the limits and interrelationships
of the genera must be based on a full consideration of the fossil forms. I do not feel myself
competent to undertake this task ; and I am aware of the fact that the conclusions drawn from
a study of the recent forms alone must be subject to a considerable amount of revision at the
hands of those who combine a knowledge of fossil and recent species.
Reptotubigera D'Orb.
Reptotubigera D'Orbigny, 1853, "Pal. Franc. Terr. Crét.", p. 751.
Stomatopora (pars), Hincks, 1880, "rust. Brit. Mar. Pol.", p. 424.
The genus Reptotubigera was introduced by D'Orbigny for forms which are entirely
adherent, having their zooecia arranged in series, as in typical species of "Idmonea". It was
regarded by him as synonymous with Obelia Lamouroux :), preoccupied by Obelia Péron and
Lesueur (Hydrozoa). Gregory 3) regards it as a synonym of Probosciua 8) Audouin.
Obelia tubulifera was described by Lamouroux from recent Mediterranean specimens,
which it might be unnecessary to separate from Ttifailipora. Proboscina boryi was named by
Audouin from specimens, obtained in Egypt, which had been figured by Savigny. The hgures
show that these are also Tiebu/ipora-like forms, with well developed peristomes, but having
ovicells of an unusually simple character. It is not stated whether they came from the Mediter-
ranean or the Red Sea coast. Most of the species figured by D'Orbigny, namely R. neocomiensis
(PI. 763, figs 1 — 3), R. marginata (PI. 750, figs 19 — 21), R. ramosa (PI. 751, figs 1 — 3), and
R. serpens (PI. 751, figs 4 — -7) are adnate forms in which the branches are linear or not much
expanded, and in which the peristomes seem to be comparatively short. With this type the
1) Lamouroux, 1821, "Exp. Méthod."', p. 81 (nee Obelia Péron and Lesueur).
z) Gregory, J. W., 1896, "Cat. Foss. Bry. Brit. Mus.", "Jurassic Bry.", p. 59.
3) For a criticism of this genus, see Walford, 1889, "Bry. Inf. Oolite Shipton Gorge", I, Q. J. Geol. Soc. XI. V, p. 565.
119
I 20
species described below agrees well ; and it seems convenient to place it in D'Orbigxy's genus.
It might well have been placed in Stoma/opora, as understood by HlNCKS; but Grkgorv (t. cit.,
p. 42) has given reasons for restricting this generic appellation to species in which the zooecia
are uniserial, as in S. granulata.
1. Reptotubigera philippsae n. sp. (PI. X, fig. 9).
Idmonea anstralis (pars), Philipps, 1899, Willey's Zool. Res., Pt. IV, "Polyzoa", pp. 441,449.
I name this species after Miss E. G. Philipps, who has recorded it from Lifu. It is
probably allied to Idmonea raniosa Waters *).
250. A. Stat. 310. 8°30'S., H9°7'.5E., 73 Metres; sand with a few pieces of dead coral.
(On an Alcyonarian axis).
and (Mus. Zool. Cambridge, Reg. Mar. 1, 1898), Idmonea australis (so determined by Miss
Philipps), Lifu, Loyalty Islands, A. WiLLEY Coll., Reg. Mar. 1, 1898.
Zoarium entirely adherent, formed by two or three elongated, strap-like lobes which
may diverge from one another at an angle of nearly 1800; the edges of the lobes forming a
basal lamina, marked by porous areas representing closed vestigial zooecia. Lobes nearly
semicircular in transverse section, most raised in the middle, but without any angulation in
that region. Zooecial series very close together, alternate in the more proximal parts, but
becoming nearly opposite at the ends of the lobes. Number of zooecia in a series 3 or 4,
less in the proximal parts of the lobes. The two or three zooecia nearest the middle of the
lobe are connate, the outermost zooecium usually separate from the others. Peristomes short,
completely connate or partially free; curved in the more proximal parts, straight and nearly
at right anp/les to the surface of the branch at the distal ends of the lobes. Ovicells infiated,
rather broad, but not very distinctly outlined. Ooeciostome a short tube, developed on the
proximal side of one of the series of zooecia, near its median end. Ooeciopore directed
frontally, elongated transversely and more or less oval.
The 'Siboga' Collection includes only two specimens of the present species; both from
the same Station. The larger, which is represented in fig. 9, measures 9 mm. in total length,
and consists of two strap-shaped lobes, closely adherent to the substratum, an Alcyonarian
axis, and diverging from one another at an angle of almost 1800. In the oldest part of the
colony the zooecia are somewhat smaller than in the more distal parts, and the full number
of zooecia in a series is not attained. Some of the zooecia near the centre have long peristomes,
which may be free. The series of zooecia are unusually close together. They are at first
regularly alternate and the zooecia are curved; but, towards the end of the lobes, where
ovicells are present, they become almost opposite; the zooecia being now straight and standing
almost at right anodes to the surface. The arrangement of the zooecia in the series has been
indicated in the diagnosis. The middle of the branch is the most elevated part, but it is
rounded and not angular.
l) Waters, A. W. 1887, "Tert. Cycl. Bry. New Zealand", Q. J. Geol. Soc, XLIII, p. 339. Miss Jei.ly (1S89, "Syn. Cat. Ree.
Mar. Bry.", p. 119) records recent specimens from Stewart Island.
120
I 2 I
Both lobes possess an ovicell near the distal extremity, and the longer lobe has a more
proximal one as well. The ovicells are all alike, rather broad and moderately swollen, but
not very distinctly outlined. The ooeciostome is in each case a very short tube, which rests
against the two inner zooecia of a series, on their proximal side. The ooeciopore is transversely
oval and is visible in the frontal view of the branch.
The basal surface of the lobes is prolonged into a well marked marginal lamina, which
projects beyond the rest of the lobe and is closely adnate to the substratum. The marginal
lamina shows a series of porous areas, which are represented in fig. 9 (/) on the right side,
and are separated from one another by low but moderately broad ridges.
The Polyzoa belonging to Dr A. Willey's Collection from Lifu and other localities
were described by Miss E. G. Philipps, and are preserved in the Universitv Museum of Zoology
at Cambridge. Two slides mounted by Miss Philipps were referred by her to I. australis\
but they certainly do not belong to MacGillivray's species. Of these, one slide contains
several young colonies, on sea-weed; showing the primitive disc, which is surrounded by fine
denticulations, like those figured by Busk r) in a form referred by him to Tttbulipora organisans
D'Orb. The marginal lamina of the specimens in question, instead of forming a broad zone round
the branch, is hardly present, but it is represented by a number of short lateral projections
which are separated from one another by rounded bays ~). The zooecia have long, slender,
curved peristomes, some of which are free and others connate. It is not impossible that these
specimens belong to the same species as the 'Siboga' specimens ; but, if so, it would have to
be assumed that the entire colonies hacl not developed beyond the condition which is indicated
in the central region of fig. 9 and that the projecting points of the marginal lamina represent
the porous areas of the 'Siboga' specimens not yet closed. In the absence of any evidence as
to the characters of the primitive disc of the 'Siboga' specimens, it woulcl, however, be unsafe
to assume that these Lifu colonies belong to the same species.
The second slide referred by Miss Philipps to I. australis may, however, be placed
without hesitation in the same species as the 'Siboga' specimens, with which it agrees closely
in the form and arrangement of the zooecia, in the ovicells (of which two are present), in
the ooeciostomes, and in the marginal lamina. The specimen throws considerable light on the
morphology of the porous areas of the marginal lamina shown in fig. 9. The agreement with
the 'Siboga' specimens is complete in the more central part: but, at the end of one of the two
lobes present, the porous areas are represented by the open mouths of structures which exactly
resemble the incompletely developed zooecia which are usually found at the growing margin of a
Cyclostome colony. It is obvious from this specimen that the porous areas result from the closure
of the imperfectly developed zooecia by calcareous plates, in which pores are developed. The ridges
separating the porous areas are clearly seen to be the remains of the lateral walls of zooecia, the
porous plates having been developed at a level slightly lower than the edges of the septal walls.
1) Busk, G., 1879, "Pol. Kerguelen", Phil. Trans., CLXV11I (Trans, of Venus Exp.) p. 6 (sep.). PI. X, figs 22 — 25.
2) It may be noted that the occurrence of fine denticulations round the primitive disc and of pointed lateral projections at the
edges of the marginal lamina cannot be regarded as distinctive of any single species, since both characters are shown in Japanese specimens
(Owstün Coll.. 40. F.), in the Cambridge Collection, the adult condition of which is of a flabellate type resembling Tubulipora jlabcllaris.
121
SIBOGA-EXTEDITIE XXVIII a. IÓ
Measurements, in n. : —
Length of the entire colony, fig. 9, measured in a straight line, 9,000 ;
Greatest diameter of right lobe, 1,330;
Width of the longer zooecial series, 360;
Longest diameter of orifices, about 100;
Width of ooeciopore, 110.
Tubulipora Lamarck.
Tubulipora Lamarck, 1816, "Hist. An. sans Vert", Ed. 1, II, p. 161.
Tubulipora Lamouroux, 1821, "Exp. Méthod.", p. 1.
Tubulipora Harmer, 1S98, "Dev. Tubulipora", Quart. J. Mier. Sci., XLI, p. 90.
Idmonea Lamouroux, 1821, "Exp. Méthod.", p. 80.
I have stated my conviction, on a former occasion1) that Idmonea, as commonly understood,
is inseparable from Tubulipora. The first species given by Lamarck is T. transversa 2), which
from his synonymy appears to be identical with the species described by most authors as
Idmonea serpens, but which appeared to me to be more properly designated T. liliacea Pall.
This species is usually adnate, although showing some tendency to become erect. In this latter
form it has so marked a similarity to species like Idmonea atlantica auctt., that it is hardly
possible to separate it generically from that species, and therefore from a number of other erect
species which writers on recent Polyzoa are agreed in referring to Idmonea. It passes, on the
other hand, through species like T. phalangea and T. flabellaris to the flabellate, adnate species
which are usually regarded as constituting the genus Tubulipora. T. phalangea shows the
serial arrangement of zooecia in as pronounced a form as in any species of Idmonea, while
T. flabellaris may have a considerable proportion of its peristomes isolated, although others
may be arranged serially. T. aperta Harm. was characterised as a species in which the peristomes
are all isolated or in which the tendency to assume a serial disposition is only slightly marked.
There seems thus to be a continuous succession, among recent species, from the erect "Idmonea"
type, with strongly marked alternating series of zooecia, to the adnate, flabellate, " Tubulipora"
type, in some species of which the serial arrangement is wanting.
While most writers on recent Polyzoa have distinguished Idmonea from Tubulipora by
its erect habit, a different view is taken by some Palaeontologists. Gregory 3) considers that
I triquetra, the only species included by Lamouroux in Idmonea, and therefore indubitably
the genotype, differs generically from "the erect forms attributed to Idmonea'. He accordingly
defines Idmonea as consisting entirely of adnate forms, and uses Crisina D'Orbigny for the
erect forms which by most authors are placed in Idmonea. For the flabellate, adnate species
referred by most authorities to Tubulipora, Gregory employs Phalangclla Gray '). I regret that
I cannot follow him in this usage, on the ground that the only two species included by Gray
1) Harmer, S. F., 1898 (see ref. on p. 124), p. 88.
2) This is the only species mentioned by Lamouroux (1821, p. 1); and, as pointed out by Gregory (1S99, "Cat. foss. Bry.
Brit. Mus.", "Cret. Bry.", I, p. 157), it thus became the genotype of Tubulipora.
3) Gregory, J. W., 1899, t. cit., p. 150.
4) Okay. J. E., 1S4S. "List Brit. An. Brit. Mus.", I, pp. 139, 149.
122
123
in his genus are P. phalangca and P. flabellaris : - - both of them species which appear to
me congeneric with Tubulipora liliacea (= T. transversa, the genotype of Tubulipord). In view
of the enormous size of the genus Tubulipora, as I understand it (including the numerous fossil
forms) some subdivision may be desirable on practical grounds; but I am unable to see how
the adnate and erect conditions can be used by themselves in subdividing the genus into genera
or subgenera.
It may be noted that Canu *) has proposed to abandon the genus Tubulipora, on the
ground that it has been used in so many different senses by various authors. For the adnate,
flabellate species referred to this genus by Hincks and others, he uses MacGillivray's genus
Liripora ~), which was, however, introduced for Diastopora lineata MacG. and D. fasciculata
MacG. : — ■ species which appear to belong to Bercnicca rather than to Tubulipora.
I have accordingly referred most of the forms of Tubuliporidae obtained by the 'Siboga'
to the genus Tubulipora. The species commonly known as Idinonea radians appears to me
to deserve generic separation; and I have placed it in Crisina D'Orbigny. For one of the
other species I have employed Reptotubigera D'Orbigny. I follow Waters in referring species
of "Idmonea" with "dorsal" ovicells to Tervia Jullien.
For the synonymy of species referred to these and other genera, Neviani's still
incomplete "Monografia del Genera fdmonea" s) should be consulted. Lang*) has published a
recent summary of the Cretaceous species of "Idmoniidae".
i. Tubulipora concinna MacG. (PI. X, fig. 10).
Tubulipora concinna MacGillivray, 1885, "Descr. new Pol.", VII, Trans. Proc. R. Soc. Vict.
XXI, p. 94, PI. I, figs 10 — \ob.
? Tubulipora lucida MacGillivray, 1885, Ibid., VIII, t. cit., p. 116, PI. V, fig. 1.
-1 co. D. j
' " I Stat. 240. Banda Anchorage, 9 — 45 Metres; black sand, coral, Litliothamnion-bank
35 '" " in 18—36 Metres.
352. B. 1
(?sp.) 57. Y. Stat. 80. Borneo Bank, 2°25'S., U7°43'E., 40 — 50 Metres; fine coral-sand.
Zoarium more or less flabellate, completely adnate. Primitive disc without denticulations.
Zooecia at the commencement of the colony strongly refiexed, the younger zooecia with
extremely long, delicate peristomes, which are mostly completely free in the centre of the
colony, but may unite in the marginal regions, where series of connate xooecia, 2 — 4 in
number, occur nearly at right angles to the edge. Ovicells of the typical Tubulipora form,
with numerous pores. Ooeciostomes free, with a more or less everted lip. Ooeciopores elongated
in one direction.
MacGillivray (1. cit., Part VII) has described three species of Tubulipora from Victoria,
under the names T. concinna, T. pulchra and T. counala, all apparently nearly related to one
1) Canu, F., 1908, "Icon. Bry. Foss. Argentine", An. Mus. Nac. Buenos Aires, XVII, p. 310.
2) See above, p. 117.
3) Neviani, A., 1900, Paite I; Pavte II, Cap. 1: 1901, Parte II, Cap. 2.
4) Lang, W. D., 1907, "Tab. view of Cret. Idmoniidae", Geol. Mag. (X. S.) Dec. V. Vol. IV, p. 122.
I23
124
another. The present species agrees well, so far as I can judge, wïth the first of these three.
In the arrangement of its zooecia, in the form of its ovicells, and particularly in the characters
of its ooeciostomes, it resembles T. aperta Harmer l), from which* it differs in the much smaller
size of the zooecia. The peristomes are slender and of great length, being mostly free in the
central parts of the colony and only united into series at the margin. The largest colony
(fig. 10) measures about 4.5 mm. in longest diameter.
The roofs of the ovicells occur at the bases of the greatly elongated peristomes, and
thus appear to lie at a low level. The outlines of the ovicells cannot be distinguished clearly
in the specimen figured ; but their number is indicated by the ooeciostomes, of which three
are present. Each is a tube of some length, standing more or less at right angles to the
roof of its ovicell, and completely free from the zooecia. The ooeciostome becomes compressed
near its free end, so that the ooeciopore is elongated. The free lip of the ooeciostome is
slightly everted.
350. D. and 351. E. were found on sea-weed; 352. 13. on what appears to be a
Chaetopod tube, perhaps a P hy llochaetopterus \ and 57. Y. on an Adcona (57. C). The last
specimen has rather coarser tubes, and more nearly approaches T. aperta in this respect. It
is possible that it does not belong to the same species as the specimens from Stat. 240, but
it has no more than the commencements of ovicells.
Measurements, in y. :
Greatest diameter of colony, fig. 10, 4,500;
Diameter of zooecia, 95 •,
Diameter of ooeciopore, 110.
2. Tuèulipora atlantica (Forbes, MSS.) Johnston. (PI. X, figs 4, 5).
Idmonea atlantica (Forbes, MSS.) Johnston, 1847, "Hist. Brit. Zooph.", Ed. 2, p. 27S, PI. XL VIII,
figs 3. 3-
Idmonea atlantica Busk, 1856, "Pol. Norway and Finmark", Ann. Mag. Nat. Hist. (2) XVIII,
p. 34, PI. I, figs 6a — e.
Idmonea atlantica Busk, 1S58, "Zoophytology", Q. J. Mier. Sci., VI, p. 128, PI. XVIII, fig. 5
(Madeira).
Idmonea atlantica (pars) Busk, 1875, "Cat. Mar. Pol. Brit. Mus.", III, p. 11.
Idmonea atlantica Hincks, 1880, "Hist. Brit. Mar. Pol.", p. 451, PI. LXV, figs 1 — 4.
Idmonea atlantica (pars) Busk, 1886, 'Challenger' Rep., L, p. 10.
Idmonea atlantica Ortmann, 1889, "Japan. Bry.", Arch. f. Naturg., Jahrg. LVI, Bd I, p. 58,
PI. IV, figs 20a, b (var. disticha Ortm.).
Idmonea atlantica Levinsen, 1894, "Zool. Danica", 4 Bd, 1 Afd., "Mosdyr", p. 76, PI. VII, figs 4, 5.
Idmonea atlantica (det. R. Kirkpatrick), Thurston, 1895, "Ramésvaram Island and Fauna G.
Manaar", Madras Gov. Mus., Buil. 3, p. 131.
Idmonea atlantica Neviani, 1900, "Monogr. Gen. Idmonea", Parte I, pp. 6, 49 (synonymy);
1901, Parte II, Cap. 2, p. 74.
Idmonea atlantica Jullien and Calvet, 1903, "Rés. Camp. Sci. Prince de Monaco", XXIII, p. 113.
Idmonea atlantica Waters, 1904, "Bry. Fr.-Josef Land", II, J. Linn. Soa, Zool., XXIX, p. 166,
PI. XXI, figs 2, 3 (the ovicell here figured is unusually short).
1) Harmer, S. I-"., 1898, "Dev. Tuiu/ipora", Quart. J. Mier. Sci., XLI, p. 101.
124
'25
Idmonea atlantica Waters, 1904, "Rés. Voy. Belgica", "Bry.", p. 90, PI. IX, fig. 5 (Antarctic).
Idmonea atlantica Nordgaard, 1906, "Bry. 2n<* Fram Exp.", Rep. 2"d Norw. Arct. Kxp., X" S,
p. 36.
Idmonea atlantica Calvet, 1907, "Bryozoaires", Exp. Sci. Travailleur et Talisman, VIII, p. 469.
Idmonea atlantica Calvet, 1909, "Exp. Antarct. Erang.", "Bry.", p. 41.
Idmonea atlantica Norman, 1909, "Pol. Madeira", J. Linn. Soc., Zool., XXX, p. 278, PI. XXXIII,
figs 1, 2.
Idmonea atlantica Thomcly, 1912, "Mar. Pol. Ind. Ocean", Trans. Linn. Soc. (2) Zool., XV,
p. 156.
Tubulipora atlantica Osburn, 1912, "Bry. Woods Hole", Buil. Bur. Fisheries, XXX, p. 217,
PI. XIX, figs 9, ga (good figures of zoarium, ovicell and ooeciostome).
Tubulipora atlantica, forma erecta Smitt, 1866, "Krit. förteckn." II, Öfv. K. Vet. Ak. Förh.
XXXIII, pp. 399, 434, PI. III, figs. 6, 7; PI. IV, figs 3 — 13 (ovicell figured).
Idmonea radians Van Beneden, 1849, "Reen. Bry. Mer du Nord" (suite), Buil. Ac. Roy. Belg.
XVI, p. 646, PI. I, figs 4—6.
The synonyms here gïven are only a small selection of the records of this species, which
is believed to be cosmopolitan in its occurrence, extending from the Arctic to the Antarctic
Ocean l).
26. A. Stat. 53. Bay of Nangamessi, Sumba, o — 56 Metres; coral-sand ; near the shore, mud.
o' -o* [ Stat. Ji- Borneo Bank, 3°27'S., ii7°36'E., 59 Metres; fine grey coral-sand.
56. A. Stat. 80. Borneo Bank, 2°25'S., II7°43'E., 40 — 50 Metres; fine coral-sand.
394. J. Stat. 144. Anchorage North of Salomakiëe (Damar) Island, 45 Metres; coral-bottom
and Lithothamnion.
353. F. Stat. 240. Banda Anchorage, 9 — 45 Metres; black sand, coral, Lithothamnion-bank in
iS — 36 Metres.
165. B. Stat. 248. Anchorage off Rumah Lusi, N. point of Tiur Island, o — 54 Metres; dredge
and reef-exploration.
371. D. Stat. 273. Anchorage off Pulu Jedan, E. coast of Aru Islands, 13 Metres; sand and shells.
(? sp.) 459. H. Stat. Si. Pulu Sebangkatan, Borneo Bank, 34 Metres; coral-bottom and Lithothamnion;
dredge and shore-exploration.
(? sp.) 394.O. Stat. 144. Anchorage North of Salomakiëe (Damar) Island, 45 Metres; coral-bottom
and Lithothamnion.
(r sp.) 414. B. Stat. 315. Anchorage North of Sailus Besar, Paternoster Islands, o — 36 Metres; coral
and Lithothamnion.
(Mus. Zool., Cambridge) Torres Straits, 131, A. C. Haddon Coll., Reg. Feb. 24, 1898.
Zoarium delicate, somewhat resembling that of Crisina radians, but the ramification
less profuse and the branches slightly reflexed, not usually becoming parallel. Basal surface
narrow and strap-like, marked off by sharp edges in the younger branches, generally slightly
concave and (in most of the 'Siboga' specimens) bearing ripple-like lines of growth, along
which the pores tend to be arranged in single transverse rows. Middle of the frontal surface
elevated ; the zooecia in alternating series, usually consisting of four zooecia, but sometimes of
five, three or even two. The median zooecium, or the first two from the middle, longer than
the outer zooecia. Ovicell a modified median zooecium, commonly beginning shortly before a
bifurcation and forking with the branch ; usually as long as many series of zooecia, in most
1) MacGii.livkay (iSS2, McCoy's 'Prodr. Zool. Vict.', Pee. VII. p. 30), in describing /. australis, suggested that it might prove
to be a form of /. atlantica. It appeavs to me quite distinct, after compaving specimens of the two foims in the British Museum.
«5
I 20
cases with a zisfzasf keel on its frontal surface. Ooeciostome in contact with a median zooecium,
which it resembles; the ooeciopore not differing much from a zooecial orifice and facing more
or less distally. The stalk of the colony becomes cylindrical by the development of a secondary
cancellated growth on its basal surface.
This delicate species has a considerable resemblance to Crisina radians in external
appearance; but differs from it in many important characters. The younger branches are
narrow and strap-like, as seen in basal view; and are without cancelli. In nearly all the
specimens referred to this species, the basal surface is marked by ripple-like lines of growth
(fig. 4). The ripple, which is slightly elevated, is convex distally, and the pores of this surface
of the branch are generally arranged in a single series along the projecting part. This character
is somewhat variable, but in several of the specimens it is an extremely pronounced feature.
The basal surface is nearly fiat, but is usually slightly concave; and it is separated from the
lateral surfaces of the branch by a sharply marked edge. In the main stalk of the colony this
arrangement becomes modified by the superposition, on the original basal surface, of a secondary
thickening, which grows in a distal direction, but does not extend far up the colony. This
thickening, which has a cancellated structure and is shown in fig. 4, corresponds with the "canaux
de renforcement" which have been described by Pergens :) in species of Idmonea.
The frontal surface of the branch is most elevated at its middle, owing to the fact that
the more median zooecia are much the most prominent. The zooecia, which are closely connate
in each series, are usually four in a series, but sometimes five or even three or two. The median
zooecium may project beyond its neighbour, and thus have a free peristome. The orifices are
nearly circular. The colony may commence with one or two alternating single zooecia; a
bi-zooecial condition next following and the full number being quickly acquired.
The ovicell constitutes a specially characteristic feature of the present species. Its length
varies with the degree of vigour in the growth of the colony. In most cases it is very long,
corresponding in length with as many as 1 2 series of zooecia of one side. Near its frontal part
it is somewhat compressed, in such a way as to form a zigzag keel, giving off a branch to
each series of zooecia, as indicated in fig. 5. Evidence that it is morphologically a modified
median zooecium is given by two facts: — (I) it commences proximally with a narrow portion
which occupies the position of a median zooecium; (II) the ooeciostome (fig. 5,0) occurs on the
median side of one of the series of zooecia in the course of the ovicell. From the fact that the
ooeciostome has a considerable resemblance to an ordinary peristome it is not very conspicuous;
and since only one ooeciostome occurs in each ovicell, the inner edge of a number of series
may have to be examined before it is discovered. The carinate condition of the ovicell is usually
well marked, but in some specimens it is hardly apparent. The ovicell generally commences
a short distance on the proximal side of a bifurcation, and forks with the branch; sometimes
bifurcating a second time when the branch divides.
The specimens 459. H. and 394. O. are rather more robust than the others. The basal
surface has rounded edges and the pores are uniformly scattered, without being confined to
1) Pergens, E., 1S90, "Rev. Bry. Ciétacé", Buil. Soc. Beige Géol., III, p. 311. text-fig. 5 (p. 312); cf. also Kirkpatrick,
1888, «Pol. Mauritius", Ann. Mag. Nat. Hist. (6) I, PI. X, fig. ■>!> [Idmonea tortuosd).
126
1 27
ripple-like lines of growth. In the absence of ovicells it is hardly possible to say what they
are; and I think it not improbable that they may belong to some other species.
414. B. is a fragment which may belong to the present species.
Measurements, in >j. :
Width of right hand branch, fig. 4, 400 ;
Breadth of the longest series of zooecia, fig. 5, 600 ;
Greatest length of the ovicell in fig. 5, measured in a straight line, (to end of left lobe) 2,800;
Diameter of zooecia, fig. 5, 100 ;
Diameter of orifices, fig. 5, up to 100.
2 a. Tubulipora atlantica (Forbes, MSS.) Johnst., var. Jïcxitosa Pourt. (PI. X, figs 1 — 3).
Idmonea flexuosa Pourtales, 1867, "Contr. Fauna Gulf Stream", Buil. Mus. Harvard, I, p. 1 1 1
(off Havana, 270 fathoms).
Idmonea flexuosa Smitt, 1872, "Flor. Bry.", I, K. Svensk. Handl., X, N" 11, p. 6, PI. II, figs 7, 8.
Idmonea atlantica, var. tennis Busk, 1875, "Cat. Mar. Pol. Brit. Mus.", III, p. 11, PI. IX
(Cf. Busk, 1886).
Idmonea atlantica, var. tennis Busk, 1886, Challenger Rep., L, p. 10.
Idmonea atlantica, var. tennis Hincks, 1880, "Fust. Brit. Mar. Pol.", p. 452.
Idmonea atlantica, var. tennis MacGillivray, 1887, "Descr. New. Pol.", XII, Trans. Proc. R.
Soc. Vict., XXIII, p. 181.
Idmonea tenclla Ortmann, 1889, "Jap. Bry.", Arch. f. Naturg., Jahrg. LVI, Bd I, p. 59, PI. III,
figs ia, 2,l>.
. ' } Stat. 105. 6°8'N., 1210 19' E., 275 Metres; coral-bottom.
477. A. )
iii.D. Stat. 156. o°29'.2S., I30°5'.3E., 469 Metres; coarse sand and broken shells.
Zoarium delicate, the branches diverging, sometimes of considerable length before
bifurcating. Basal surface narrow, without sharply marked edges, the longitudinal septal lines
very distinct, but without transverse ridges. Branches sub-triangular in transverse section, the
median zooecia the most prominent. Zooecial series usually with two zooecia, of which the
median one is the longer, often considerably longer than the other. The zooecia may be
completely connate, but both may develop a free peristome. Ovicells short, dilating gradually
from their narrow pro.ximal portion, then becoming considerably inflated, but without ridges on
the frontal surface. Ooeciostomes ?
The longest fragment (iii.D.) is only 15 mm. in length. This specimen and 80. A.
have branches which reach a length of about S mm. before bifurcating, but the unbranched
regions of 477. A. are shorter. The whole colony is very delicate, and the branches are narrow
and without sharp edges separating the basal from the lateral surfaces. The basal surface is
somewhat convex, and it appears to be characteristic of this form that the longitudinal septal
lines are here very distinct, and that the "back" is smooth and without ridges. The lateral
surfaces slope fairly steeply to the middle of the frontal surface. The series are bi-zooecial in
most cases, though three zooecia occur rarely. The median zooecium is usually distinctly longer
than the outer one.
127
i2;
The ovicell is not unlike that of a typical Crisia in its general proportions, and in the
fact that the narrow proximal part expands gradually into the inflated part. In length the ovicell
corresponds with 21/,, to 3 zooecial series of one side. It usually occurs at about the middle of
the unbranched part of a branch, but it may be found just on the proximal side of a bifurcation.
Other ovicells may be developed on the distal side of the first one. Thus in iii.D. the fertile
part of the branch bifurcates in one instance, and of the two branches thus formed one has
an ovicell in the middle of its length, and the other has formed a new bifurcation, one of the
branches of which bears another ovicell. In all cases observed the ovicell is simple, not bifurcating
with the branch.
Most of the ovicells are either incomplete or broken; and I have not been able to
distinguish the ooeciostome with certainty. In fig. 1 it is perhaps the second opening from the
middle of the series of four tubes which constitute the second zooecial series from the proximal
end, on the right side of the figure.
Two specimens from Japan, in the Collection of the University Museum of Zoology at
Cambridge, show considerable resemblance to the present form in their zooecial characters. But
there are differences in the ovicells, particularly in the existence of a long ooeciostome projecting
laterally on one side of the ovicell and curving distally, which make it doubtful whether they
should be referred to the same species or not.
M easu remen t s , in a:
Width of branch, fig. 2, 300 ;
Width of branch, just before bifurcation, fig. 3, 500 ;
Length of ovicell, fig. 1, 1,300;
Diameter of orifices, about 100.
The specimens here considered are both from deep water (275 — 469 Metres), and may
represent an abyssal form of the species from shallower water which I have referred to T.
atlantica. The description, unaccompanied by a figure, which was given by Pourtales would
have been insufficiënt by itself for identification-, but the specimens figured by Smitt, who
regarded I. Jïexnosa as a form of /. atlantica, have a close resemblance to those dredged by
the 'Siboga'. Busk's var. tennis of the same species seems indistinguishable from Smitt's form.
In the biserial character of their zooecia the 'Siboga' specimens resemble I. gracillima Busk ')
(nee Ortmann). Busk states that the outer zooecia are the longer; but, from the examination
of the slide labelled "Adventure Bank, Porcupine, 92 fathoms", in the British Museum I do
not feel convinced that this is correct. The specimen consists of two minute fragments which
are probably more or less worn. The median zooecia are much more distant from the basal
surface than the outer zooecia, as in T. atlantica. It is to this assemblage of species, in which
the median zooecia are at the same time the most raised and the longest, that Busk's specimen
seems to me to belong-.
From the locality in which they were found, the 'Siboga' specimens might be supposed
1) Busk, G., 1875. p. 14. PI. VII, figs 5. 6. I. gracillima Busk is antedated by /. gracillima Retiss, 1S69, "Pal. Stud. iilt.
Tertiarsch. Alpen", II. Denkschr. Ak. Wiss. Wien, math.-naturw. Cl. XXIX, p. 2S2.
128
i 29
to be identical with /. tuberosa D'Orbigny v) from Basilan Island (Philippines). But this species,
although delicate and with biserial zooecia, is said to have the series "peu distinctes", and the
two zooecia in each series are "écartées". It is further described as being everywhere wrinkled.
T. tcnella was described by Ortmann, from Sagami Bay, Japan, 70 — 100 fathoms. The
branches are described as very thin ("fadendünn"). The zooecia are usually 2 — 3-serial, most
often biserial. Ovicells are not mentioned. No measurements are given in the text, but fig. 3^
is said to be X IO- This gives a diameter of the branches, between the series of zooecia, of
600 u.; which is twice as much as the width of the 'Siboga' specimen indicated above.
3. Tubulipora pulckerrima Kirkp. (PI. IX, figs 1 — 5).
Idmonea pulcherrima Kirkpatrick, 1S90, "Hydr. Pol. China Sea", Arm. Mag. Nat. Hist. (6), V,
p. 22, PI. IV, figs 6—6/7 (Tizard Reef, 6 fathoms).
Idmonea interjuncta Waters, 1887, "Bry. N. S. Wales", I, II, III, Ibid., (5) XX, PI. VI, fig. 29,
- PP- 84, 256.
Idmonea interjuncta Philipps, 1899, WlLLEV's "Zool. Res.", Pt 4, "Polyzoa", pp. 441, 449
(Lifu, Loyalty Islands).
Idmonea interjuncta Waters, 19 14, "Mar. Fauna Brit. E. Afr.". "Bry. Cycl.", Proc. Zool.
Soc, p. 846, PI. II, fig. 5.
? Idmonea milneana Haswell, 1880, "Pol. Queensland", Proc. Linn. Soc. N. S. Wales, V, p. 35.
Idmonea milneana (pars), Neviani, 1900, " Monogr. Gen. Idmonea'', Parte I, p. 23.
Idmonea milneana Thornely, 1905, HERDMAN's "Rep. Pearl Oyster Fisheries", Suppl. Rep.
XXVI, Polyzoa, p. 127 (Ceylon).
't Idmonea radicata Kirkpatrick, 1888, "Pol. Mauritius", Ann. Mag. Nat. Hist. (6) I, p. 83,
PI. IX, figs 2, 2 a.
449. A. Stat. 59. Western Entrance Samau Strait, 390 Metres; coarse coral-sand with small
stones.
260. D. {„ \ Haingsisi, Samau Island, Timor, o — 16 Metres; Lithothamnion.
y ( Stat. 303 ) ö °
47. A. Stat. /j. Borneo Bank, 30 27 'S., II7°36'E., 59 Metres; fine grey coral-sand.
57. N. Stat. 80. Borneo Bank, 2°25'S., H7°43'E., 40 — 50 Metres; fine coral-sand.
(? sp.) 62. M. Stat. 99. Anchorage off North Ubian, 16 — 23 Metres; Lithothamnion-bottom.
S4. A. Stat. 105. 6°8'N., I21°I9 E., 275 Metres; coral-bottom.
482. B. Stat. 116. West of Kwandang Bay, entrance, 72 Metres (chart); fine sand with mud.
92. H. Stat. 117. Kwandang Bay, entrance, 80 Metres (chart); sand and coral.
99. A. Stat. 129. Anchorage off Kawio and Kamboling Islands, Karkaralong group, 23 — 31
Metres; sand.
108. L. Stat. 144. Anchorage North of Salomakiëe (Damar) Island, 45 Metres; coral-bottom
and Lithothamnion.
505. A. Stat. 164. i°42'.5 S., I30°47'.S E. 32 Metres; sand, small stones and shells.
350. C. ) Stat. 240. Banda Anchorage, 9 — 45 Metres; black sand, coral, Lithothamnion-bank
352. A.J in iS — 36 Metres.
425. A. Stat. ? , Banda Sea (on sea-weed).
165. A. Stat. 24S. Anchorage off Rumah Lusi, N. point of Tiur Island, O — 54 Metres.
276. D. Stat. 250. Anchorage off Kilsuin, W. coast of Kur Island, 20 — 45 Metres; coral and
Lithothamnion.
' '[ Stat. 251. 5°28.4S., i32°o'.2E., 204 Metres; hard coral-sand.
1) D'Orbignv, A., 1853, "Pal. Frang. Teir. Crct.", V. p. 732.
129
SIBOGA-EXFEDITIE XXVIII a. 17
\--. I. Stat. 257. In Du-roa Strait, Kei Islands, o — 52 Metres; coral.
20S. F. Stat. 282. Anchorage between Nusa Besi and the N. E. point of Timor, 27 — 54 Metres;
sand, coral and Lithothamnion.
223. A. Stat. 305. Mid-channel in Solor Strait, off Kampong Menanga, 1 13 Metres; stony bottom.
412. A. Stat. 310. 8° 30' S., M9°7".5E., 73 Metres; sand with a few pieces of dead coral.
Also, in the Collection of the University Museum of Zoology, Cambridge : —
Torres Straks, 194. A. C. Haddon Coll., Reg. Feb. 24, 1898.
Singapore, 5 — 10 fathoms, F. P. Bedford Coll., Reg. Nov. 11, 1899.
Ceylon, L. R. THORNELY Coll., Reg. Apr. 25, 1906 (determined by Miss ThüRNELY as
Idmonea milneana).
Lifu, Loyalty Islands, A. WlLLEV Coll., Reg. Mar. 1, 1898 (determined by Miss PHILIPPS as
I. interjuncta).
Port Denison, Queensland, Miss E. C. JELLV, Reg. May 24, 1895.
Japan, Uraga Channel, off Tokyo, 30 fathoms, A. OWSTON Coll., 49. K, 7. S + 15. B, Reg.
June 23, 1902.
Zoarium large, reaching a diameter of at least 45 mm. (412. A.); its branches commonly
in one layer, often nearly in one plane, but sometimes variously curved. In the majority of
cases the zoarium is more or less reticulate, owing to the existence of cross-connexions between
the branches. Stout rooting processes, consisting of a bundie of parallel kenozooecia *), are
usually given off, here and there, from the middle of the basal surface of the branches. Basal
surface not limited by sharp edges, flat or gently convex, the longitudinal septal lines distinct;
usually without transverse lines of growth. Frontal surface of branch nearly fiat, without a
median ridge. Series consisting of 2 — 3 zooecia, rarely 4, the outer zooecia the longest. Some
of the median peristomes are commonly somewhat separated from their neighbours of the same
series, and appear more or less isolated in the middle of the branch. Ovicell variable, either
simple or bifurcating with the branch, the frontal surface only slightly convex. Ooeciostome
typically single, with a much infiated basal portion, of an urndike shape. Accessory ooeciostomes
are rarely present, and may be of simpler structure than the principal ooeciostome.
This species is represented by a number of specimens in the 'Siboga' dredgings. As
in the case of so many other Cyclostomes, there is a considerable amount of variation in
size, arrangement of zooecia and other characters. These variations may depend on the general
vigour of growth. Thus the ovicells shown in hg. 4 are developed in the course of branches
of a large reticulate colony; while in fig. 1 the ovicell is formed in a very small colony whose
zooecia had not attained the typical adult arrangement at the time when the ovicell commenced
to develop.
In well branched colonies the zooecia have the disposition shown in those parts of
fig. 4 where an ovicell is not present. The rather flat frontal surface of the branch is associated
with the fact that the median zooecia are not specially raised. The alternating series consist
most commonly of two or three zooecia, or alternately of two and three; the outer zooecia
being the longest and often having their peristomes much prolonged. The median peristomes
are frequent!)- separated from their next neighbours by a distinct interval, as shown in the
1) This term was introduced by Ltsvinsen (1902, "Studies on Bryozoa", Vidensk. Mcdd. Naturb. Foren. Kjobenhavn, p. 3) for
zooecia "which lack both a polypidu and an orifice".
130
'3i
lower part of the left hand branch in fig. 4; and thus appear as isolated, nearly vertical units
which lie in the middle of the branch. In the older parts of the colonies each "series" may
be reduced to a single zooecium, as shown in the proximal part of fig. 1 ; the normal disposition
being attained in younger parts of the same branch. Minute calcareous teeth commonly
project into the peristomes.
One of the most characteristic features of the 'Siboga' specimens is the presence of
cross-connexions between the branches, as shown in figs 4 and 5. These vary greatly in their
degree of development, even in different parts of the same colony. Thus in specimen 99. A.,
from which the figures just referred to are taken, the following conditions were observed:
(I) A tube (kenozooecium) which has all the relations of the outermost zooecium of its
series grows across to the next branch, with some part of which it unites firmly, leaving a
suture between itself and the branch with which it has united (figs 4, 5). The kenozooecium
develops no orifice;
(II) the connexions may be widely separated from one another, giving rise to elongated
meshes, or tvvo may develop close together; sometimes, as in fig. 5, one from each of the
participating branches ;
(III) the connexion may be two zooecia wide, and orifices may develop on the zooecia
which take part in its formation (fig. 4, middle of figure) ;
(IV) a stouter connexion may be formed, by a bundle of several kenozooecia which
develop no orifices ;
(V) if the next branch does not lie in a suitable position the connexion may run across
it, without meeting it, and may unite with the branch next but one to that from which it has
taken origin.
In addition to these formations a considerable proportion of the specimens develop long,
straight, rooting columns l) (fig. 3), formed of a bundle of as many as 8 kenozooecia, which
diverge when they reach the substratum, sometimes separating from one another to form
irregular attaching processes. These may give rise to isolated, free peristomes, which reach a
considerable length (350. O2).
The largest specimen found (412. A.) consists of a colony about 45 mm. in diameter. It
is completely overgrown by, and embedded in a Sponge, so that its form cannot be made out.
On isolating one or two branches of this colony by means of Eau de Javelle, it was found that
the peristomes were of unusual length, and that they were to a considerable extent discrete ; —
probably as a result of an attempt to keep pace with the growth of the including Sponge.
The ovicells are generally about as long as three series of zooecia of each side (fig. 4);
their frontal surface being only slightly convex. In some colonies, as shown in the figure
referred to, they are "simple"; i. e., not extending into a bifurcation. The ovicell appears to be
a modification of a median zooecium. Its ooecistome, when typically developed, is a beautiful
urn-shaped structure, with its basal region much dilated and porous; then contracting slightly,
1) As in Idmonea radicata Kirkpatiick (1888, p. 83, PI. IX, fig. 2), with which this species may be identical. Similar rooting
columns have also been described, in /. pedata^ by Norman (1909, "Pol. Madeira", J. I.inn. Soc, Zool., XXX. p. 279. PI. XXXIII, figs 6, 8).
131
132
and fïnally expanding to form an everted lip which surroünds the ooeciopore. In the majority
of cases the ooeciopore is transversely elongated and looks frontally and distally; but rarely
it faces in a proximal direction. It is usually symmetrically situated, near the middle line of
the ovicell.
The development of the lip of the ooeciostome is variable. In fig. i it is wider than in
fig. 4; and in some cases it is wider still. In other colonies, on the contrary, the lip is hardly,
or only slightly developed; and even the basal dilatation may be absent. But the same colony
may show one or two ooeciostomes of the typical form, even thongh other ovicells have this
structure in what may be considered the reduced form above described.
In nearly all cases, each ovicell has a single ooeciostome, even when it bifurcates with
the branch. But in specimen 350. C, where the ovicell may bifurcate once or even twice,
several accessory ooeciostomes are present. One or more of these. may have the typical form;
but they are usually simpler in character. A case of this kind is represented in fig. 2, where
the ovicell has two main branches, each of which shows the indication of a second bifurcation.
Each of the principal branches has three ooeciostomes (t>) facing in various directions. The most
proximal ooeciostome in the branch of the left side is more or less of the typical form, although
not fully developed and without an everted lip-, and this is perhaps the normal ooeciostome
of the ovicell. The other five are less like what I have described as the typical ooeciostome
of the species.
The form described by Kirkpatrick as Idmonca pulcherrima (see synonomy) agrees
closely with 'Siboga' specimens (e. g. 99. A.), as I have convinced myself by an examination
of the type-specimen (89. 8. 21. y^i) m tne British Museum, in the form of the colony, in its
reticulate arrangement, in its zooecial characters, and in its ovicells and ooeciostomes. The
description given by that author makes no reference to the occurrence of more than one
ooeciostome in an ovicell; but his fig. 6a shows an ovicell with two ooeciostomes of the typical
form. I have previously called attention to the occurrence of accessory ooeciostomes in Tubu-
lipora aperta 1), from Norway.
The polypides have the loop of their alimentary canal unbent during retraction : — a
character which seems to be general in Cyclostomes,. and is probably associated with the long
and narrow shape of their zooecia. The distal ends of the tentacles include a considerable
number of the "excretory vesicles" which I have described in other species of Tubulipora~)\
and similar structures appear to occur in the embryos. The retractor muscles diverge from a
single point on the wall of the zooecium, which is situated at some distance on the distal side
of the apex of the caecum of the stomach (in the retracted polypide).
The primitive disc of the colony has no marginal denticulations.
The present species appears to be closely related to T. milneana D'Orb. 3) and T.
interjuncta MacGill. 4), if indeed it is not identical with one or both of those forms. If these
1) Hakmeu, S. F., 189S, "Dev. Tubulipora\ Quart. J. Mier. Sci., XI.I, pp. 101, 104. PI. VIII, fig. 2.
2) t. cit., p. 113.
3) D'Orbigny, A., 1S46, "Voy. Am. Mérid.", V, 4e Partie, "Zoophytes", p. 20 (as Idmonea milneana\ PI. IX, figs 17—21.
4) MacGii.uyray, 1'. II., 1SS6, uDescr. new Pol", IX, Tr. Pr. R. Soc. Vict., XXII, p. 137 (as Idmonea interjuncta).
132
species are distinct from one another it is probable that they have often been confused by
other writers.
D'Orbignv's specimens were obtained in the Falkland Islands, and his description and
figures represent a form which is much coarser in its habit than the 'Siboga' specimens, the
series consisting of 3 — 4 zooecia. The basal surface of the branches appears to be very convex ;
and there is no indication of cross-connexions or of rooting columns l). A specimen obtained
by Mr Darwin, during the 'Beagle' Voyage, at the Albrolhos Islands, Brazil, and now in the
Collection of the University Museum of Zoology at Cambridge, agrees well with D'Orbignv's
account. The branches are robust, the zooecia being usually 4 in each of the alternating series.
When an ovicell is formed the branch widens greatly, and the number of zooecia in one series
rises to 6 or 7 ; represented, as in the 'Siboga' specimens, by peristomes isolated from one
another by the roof of the ovicell. A similar enlargement of the branch may occur at a
bifurcation, without the formation of an ovicell. The only ooeciostome which is satisfactorily
seen is much compressed, with a narrow ooeciopore, much elongated in one direction, looking
directly frontally and placed obliquely. The much coarser habit, with the absence of connexions
between the branches, together with the characters of the zooecial series and of the ovicell,
appear to me to point to the fact that the 'Siboga' specimens do not belong to D'Orbignv's
species. The basal surface of Mr Darwin's specimen is very broad, nearly flat in the middle,
and shows distinct longitudinal septal lines and a certain number of transverse lines of growth.
T. interjuncta was described, but not figured, by MacGillivray from Victorian specimens.
In the existence of connexions between the branches and of rooting columns from their basal
surfaces, it obviously comes very near the Malay specimens. But although MacGilliyrav
describes it as resembling a slender form of T. milneana, the evidence of Victorian specimens
in the Cambridge Collection shows that it is a more robust form than those from the 'Siboga'
clredgings. The Victorian material alluded to includes only a single ovicell; which possesses a
marked rounded swelling in the middle line. On the proximal side, this swelling opens by an
ooeciostome which is hardly represented by more than a greatly compressed, slit-like ooeciopore,
opening almost directly from the swelling in a vertical plane, and facing proximally. The long
axis of the ooeciopore is transverse, and is convex proximally, the slit being vertical and
accordingly only visible when the ooeciostome is looked at from the proximal end of the branch.
The zooecia are usually 3 — 4 in a series, and are distinctly larger than those of the 'Siboga'
specimens. I do not feel convinced that these are identical with MacGillivray's species.
Waters2) has compared T. milneana and T. interjuncta from Port Jackson; but I do
not feel sure that his determination of T. interjuncta is correct. The characters of the ooeciostome
in Waters' specimens appear to show that these belong to the same species as the 'Siboga'
material; and that they are hence not identical with MacGillivray's species. The ovicell of a
Mediterranean specimen (Capri) which has been described as I. milneana and figured by Waters8)
1) These structures may, however, occur in this species (Waters, A. W., 1905, Tin'. Cape Hovn", J. Linn. Soc, Zool.,
XXIX, p. 249).
2) Waters, A. W., 1SS7, "Bry. N. S. Wales", Pts II, III, Ann. Mag. Nat. Hist. (5) XX. PI. VI, fig. 29: pp. 256, 257.
3) Ibid., 1S89, '•Ovicells Cycl.", J. Linn. Soc, XX, p. 279, Tl. XIV, Hg. 8.
133
134
has not much bearing on the question at issue if, as I think, the determination of the species
is uncertain.
It may be admitted that the synonymy of this species is doubtful. I have preferred to
describe the 'Siboga' specimens as T. pulckerrima, as I am quite satisfied that they belong
to the species so named by Kirkfatrick in 1890. But it is not impossible that it should
with more propriety be referred to T. interjuncta MacGillivray, 1S86 or to T. radicata
Kirkpatrick, 1S88.
The following notes may be added with regard to the specimens in the Cambridge
Collection which are considered to belong to the present species : —
Torres Straits. Ovicells are not present. The zooecia are robust, and the
branches show no cross-connexions. Basal rooting columns are present.
Singapore. — A very typical specimen, with cross-connexions and rooting columns.
Ovicells, resembling those shown in fig. 4, are present, but they have produced no ooeciostomes.
It may be inferred that development of the embryos had not proceeded normally.
C e y 1 o n (determined by Miss Thornely as I. mi/neana). — Two small fragments
with ovicells. The ooeciostomes, of which two are visible, do not show the urn-like shape of
those of the 'Siboga' Collection. A basal rooting column is present, but no cross-connexions
are visible.
Lifu, Loyalty Is (determined by Miss Philipps as I. interjuncta). — 4 slides, one
of which shows ovicells and typical cross-connexions. The ooeciostomes are intermediate in
character between those of the 'Siboga' specimens and those from Ceylon. Basal rooting
columns occur.
Oueensland. — Fragments, without cross-connexions, rooting columns or ovicells,
but probably belonging to the present species.
Japan, Owston Collection (49. K., 7. S. + 15. B.). — Fragments, with ovicells showing
urn-shaped ooeciostomes precisely like those of the 'Siboga' specimens. Rooting columns are
present, but no cross-connexions.
Measurements, in u. : —
Diameter of left branch, fig. 4, on the proximal side of the ovicell, 700 ;
Diameter of left branch, fig. 4, on the distal side of the same ovicell, 500 ;
Diameter of the distal end of the same ovicell, 1,100;
Length of its inflated part, 2,000;
Diameter of the inflated part of its ooeciostome, 210;
Diameter of the ooeciopore, 255;
Diameter of the zooecia, 180 — 200 ;
Diameter of the orifices, about 1 So ;
Greatest width of the branch, between the zooecia, fig. 5, 1,000 ;
Greatest width of the ovicell, fig. 2, 3,200;
Width of the ooeciopore, fig. 1, 280 ;
Width of the rooting column, near its origin, fig. 3, 220;
Length of the same rooting column, 3,400;
134
i35
4. Tubulipora cassiformis1) n. sp. (PI. IX, figs 6 — io).
Type. 94. A. Stat. 119. i°33'.5N., i24°4i'E., 1901 Metres; stony bottom.
112. A. Stat. 156. o°29'.2S., i30°s'.3E., 469 Metres; coarse sand and broken shells.
Zoarium reticulate, with the branches arranged in one plane. Strong rooting columns,
formed of a bundle of kenozooecia given off occasionally from the basal side of the colony.
The branches are commonly parallel to one another, and the cross-connexions are often arranged
at the same level in adjacent branches, thus giving rise to elongated, narrow meshes of uniform
size. In other parts the meshes may be more irregular. Basal surface smooth, porous, the
zooecial outlines indistinct. The branch is subtriquetrous in transverse section, the middle of
the frontal surface being the part which is most prominent. This portion is occupied by an
alternating series of zooecia, the peristomes of which are somewhat compressed and bend
outwards, the orifice being oval and the part of the distal end of the zooecium which faces
outwards being flattened. Each "series" of zooecia is thus reduced to a single member,
although the outer members are represented by kenozooecia which form the lateral parts of
the branch and from which the transverse connexions are developed. Ovicells elongated, usually
corresponding in length with about three zooecial series on each side; considerably inflated,
the swollen part beginning suddenly at the proximal end. The ovicell encircles the median
zooecia on their outer sides, and is here produced into an angular ridge. Ooeciostome short,
the ooeciopore being a wide aperture, near the distal end of the ovicell.
This very characteristic deep-water species is represented in two of the 'Siboga' dredgings,
in each case by a small amount of material. The specimens arrived broken, so that it is not
possible to decide what was the form of the complete colony. The longest fragment measured
about 19 mm. in length. The species is distinguished by its reticulate character, by the reduction
of each "series" to a single complete median zooecium and an outer kenozooecium, and by
the narrow inflated ovicell.
The fragments are all flat, the branches being arranged in a single plane. Those from
Stat. 119 have elongated meshes, with parallel sides, as shown in fig. 8, a basal view. The
connexions between adjacent branches are formed by subcylindrical bundies of kenozooecia,
several of which constitute each bundle. In this specimen most of the connexions are developed
at the same level in neighbouring branches, so that a very regular disposition of the meshes
results. The connexions may be quite transverse, or somewhat oblique; and they are formed
from one or both branches. Both these conditions are shown in fig. 8 ; the region where the
bundle of kenozooecia meets the other branch or a similar bundle developed from it being
distinctly indicated by a suture. Since each bundle dilates as it approaches the suture, a
transverse or oblique ridge is formed where two bundies unite. The connecting bundies may
have a compressed shape, as shown in fig. 10. A massive column of kenozooecia may be given
off by the basal surface of a branch, and 110 doubt indicates an attaching arrangement.
The specimens from Stat. 156 may have belonged to two colonies, as there are well
1) From cassis, a net ov cobweb.
135
136
marked differences between them. One of them consists of branches of the type described in
the other specimen, except that the meshes are more irregular in size. The other has branches
which are rather more divergent and less parallel. In this specimen the zooecia agree with
those of the colony from Stat. 119; but in the other piece, which has parallel branches, the
zooecia exhibit the curious modification which is illustrated by figs 6 and 9. The median part
of the wall of the peristome is here prolonged into a flat, shield-like portion which almost or
entirely conceals the orifice, in a frontal view of the branch. In a side view (fig. 9) it is seen
that the terminal portion of the peristome comes off at right angles from the shield-like part,
which is prolonged beyond it. The same specimen has the lateral edges of the basal wall of
its branches distinctly serrate (figs 6, 9, 10), the projecting parts being transverse and joining
the rest of the branch at right angles distally, and sloping gradually in the proximal direction
into the side of the branch. The occurrence of septal lines outlining these serrated portions
(fig. 9) indicates that the parts in question are really kenozooecia and are thus outer zooecia
which develop no orifices. It may be concluded that the zooecial series consist of two zooecia,
of which the median one possesses an orifice and is a complete zooecium, while the outer one is
reduced to a kenozooecium. This serrated condition has not been found in the other specimens,
in which, however, the lateral parts of the branches appear to be similarly constituted of
kenozooecia, which do not project in the way just described. Since the transverse connexions
between the branches are produced from these lateral regions, it may be concluded that some
of the lateral zooecia have been modified into kenozooecia which unite in bundies to form the
cross-connexions. These facts seem to indicate that the present species is comparable with other
species of Tubulipora in which several zooecia unite laterally to form the alternating series.
The complete zooecia always have the outer parts of the distal portion flattened.
The ovicells are "simple", in the sense that they are dilatations which do not bifurcate
with the branch. Their form is shown in fig. 7, where it will be seen that the ovicell, although
much infiated, is not much wider than the branch. The inflated part of the ovicell commences
abruptly at its proximal end. The ovicell extends round the zooecia which occur in its region,
on their outer sides; and an angular ridge is formed at the side of it, distally to each complete
zooecium, and sometimes elsewhere. In all the specimens in which an ooeciostome occurs, this
structure is found at the distal end of the ovicell, on one side. It appears as a wide tube, the
ooeciopore of which looks frontally (see the right side of the distal end of the ovicell in fig. 7) ;
but I am not sure that the ooeciostome is fully formed in any of the ovicells present.
M e a s u r e m e n t s , in u. : —
Width of branches, between the cross-connexions, fig. 8, 340 — 410 ;
Diameter of cross-connexions, fig. 8, 190 — 210;
Length of infiated part of ovicell, fig. 7, 1,700 ;
Greatest width of proximal part of ovicell, fig. 7, 635;
Greatest diameter of orifices, up to 175;
Diameter of zooecia, fig. 6, 150.
136
'37
Crisina D*Orbigny.
Crisina D'Orbigny, 1850, "Prodr. Pal. Stratigr.", II, p. 265 (nomen nu duin).
Crisina D'Orbigny, 1853, "Pal. Frang. Terr. Crét.", V, pp. 912, 728.
Crisina Neviani, 1900, "Monogr. Gen. Idmonea", Parte II, Cap. 1, p. 65.
(nee Crisina Gregory, 1899, "Cat. Foss. Bry. Brit. Mus.", "Cret. Bry.", I, p. 159.)
Crisina, as I use it in this Report, differs from the erect species of Tubu/ipora {Idmonea
auctt.) in the development of cancelli, specially in longitudinal lines along the basal surfaces of
the branches. These cancelli are developed in the earliest stages of the growth of the branch in
the recent C. radiaus, the ovicells of which differ from those of typical "Idmonea" in possessing
curious lateral porous windows.
The history of the generic names Crisina and Crisisina has been given by Gregory
(t. cit., p. 159, note) ; but I do not find myself able to agree with him in his conclusions.
Gregory considers that D'Orbigny used Crisina and Crisisina indiscriminately, and that the
employment of the latter term was probably due to a misprint. I think, on the contrary, that
there is clear evidence that D'Orbigny meant to distinguish the two genera, and that there is
reason for accepting Crisina as he defined it.
In the "Prodrome", Vol. II, p. 175, D'Orbigny defines Crisisina as follows : — "C'est
une Idmonea, dont les branches sont libres au lieu d'ètre fixes ". Crisina occurs on p. 265 of
the same volume, five species being mentioned. Xo diagnosis is given; and Crisina thus appears
to be a nomen nudum, so far as this publication is concerned. Gregory is perfectly
correct in stating that the species of both genera occur under Crisisina in the Index of the work
(Vol. III, 1852, p. 51); but a careful examination of this index shows that there are several
other mistakes in it, so that too much stress need not be laid on the omission of Crisina. That
D'Orbigny intended, even in this work, to keep the two genera distinct is indicated by the fact
that Crisisina and Crisina appear with two separate generic headings on p. 265 of Vol. II.
In his larger work ("Pal. Frang.", p. 728) D'Orbigny states that he had given the name
Crisina to the species with erect branches, when he would have been more accurate in stating
that Crisisina was the name referred to ; while on Plate 612 he uses Crisisina in one place
where the text shows that he meant Crisina. But his real meaning may be inferred by noticing
that in this work he entirely suppresses Crisisina, placing species referred to it in the smaller
work under Idmonea, and stating specifically (p. 728) that he now places species with free
branches and without inferior pores in Idmonea, and that he retains Crisina for species "dont
la face inférieure est criblée de pores spéciaux".
A comparison of D'Orbigny's two works with regard to the treatment of the five species of
Crisina mentioned in the "Prodrome" confirms the above conclusions. The facts are as follows : —
"Prodrome", Vol. II. "Pal. Frang. Terr. Crét.", Vol. V.
Crisina unipora p. 265. Idmonea unifora p. 737, PI. 613, figs 1 — 10.
Crisina rainosa p. 266. Idmonea ramosa p. J36, PI. 611, figs 11 — 15.
Crisina Normaniana p. 265. Crisina Normaniana p. 914, PI. 612, figs. 1 — 5.
Crisina sub gr adat a p. 266. Crisina subgradata p. 914, PI. 612, figs 6 — 10.
Crisina triangularis p. 266. Crisina triangularis p. 915, PI. 612, figs ii — 15; PI. 614, figs 11 — 15;
PI. 769, figs u — 14.
137
SIBOGA-F.XrEDITIE XXVIII a. iS
138
The names on the Plates do not, however, agree completely with those used in the text.
It should be noted that the three species retained in Crisina in the "Paleontologie
Frangaise" have porous backs, and thus agree with the definition of Crisina given in that work.
Of the others (as shown by the figures) I. unipora has no pores on its basal surface; and it
is stated in the text (p. 736) that the appearance of pores in that position in I. ramosa is due
to the fact that the specimens are worn.
Gregory further objects (p. 160) that on PI. 614 of the larger work D'Orbigny figures
three species marked as Crisina, and that of these C. cenomana shows pores on the basal
surface. But D'Orbigny expressly states in his text (p. 733) that the appearance of these pores
is due to wear. C. cenomana (p. 732) was accordingly placed by D'Orbigny in Idmonea, to
which genus the second species, C. snbgracilis, a form with an imperforate back (p. 738), was
also referred. The only species left on this Plate is there called Crisina Ligeriensis, which is
treated in the text as a synonym of C. triangularis . Gregory's statement that C. cenomana is
"the only species amongst those originally included in Crisina which is available as a member
"of that genus" is invalidated by the fact that in the "Prodrome" (Vol. II, p. 175) C. cenomana
is referred to Crisisina, and not to Crisina. I have indicated above that Crisina as used in
the "Prodrome" appears to be a nomen nudum; but even if its validity in that work is
admitted, it may be pointed out that three of the five species originally included in it are
retained by D'Orbigny in Crisina in his larger work; in which, in fact, he rejects Crisisina
but retains Crisina.
Gregory uses Crisina (p. 160) for the erect species which most writers on recent Polyzoa
refer to Idmonea. The above considerations appear to show that this procedure is inadmissible;
though it would have been open to him to have used Crisisina in the sense in which he
actually used Crisina.
The peculiarities of the recent "Idmonea" radians appear to be sufficiently marked to
justify its separation from most of the recent species of " Idmonea\ and as it agrees exactly, in
its porous basal surface, with the feature emphasised by D'Orbigny in defining Crisina, there
seems to be every reason for placing it in that genus. The basal surface of C. radians has
the closest resemblance to that of C. triangularis, as shown by D'Orbigny in his PI. 769,
fig. 13. As I show below, in my account of C. radians, this feature is pronounced from the
earliest stages in the °Towth of the branches, and thus differs from the arrangement found in
other species here described (cf. Tubulipora atlantica, p. 126), in which the stalk of the colony
becomes thickened by secondarily developed calcareous tubes, the "canaux de renforcement"
of Pergens.
I cannot find sufficiënt evidence of the characters of the ovicells of fossil species which
fall within Crisina in D'Orbigny's sense. MacGillivray l) has described the ovicell of a Tertiary
Victorian form referred by him to C hocJistcitcriana Stoliczka, as being dorsal in position.
Waters 2), who regards Stoliczka's species as identical with C. radians, considers that the
form described by MacGillivray is Horncra fissnrata Busk. I am not quite convinced that this
1) MacGillivray, P. IL, 1895, "Monogr. Tert. Pol. Vict.", Trans. R. Soc. Vict. IV, p. 121, PI. XVI, fig. 14.
2) WATERS, A. W., 1914, "Mar. Fauna Brit. E. Afr.", "Biy. Cycl.", 1'roc. Zool. Soc, p. 845.
138
139
latter identification is certain ; since MacGilliyray's figures, both of the unfertile branches and
of the ovicell, show a considerable resemblance to C. radians. It is just possible, therefore,
that Crisina may have ovicells of the C. radians type indifferently on the frontal or the basal
surface of the branch .
i. Crisina radians Lamk. (PI. X, iïgs 6 — 8).
Retepora radians Lamk, 1816, "Hist. Nat. An. s. Vert.", II, p. i S3 (Nouvelle-Hollande).
Retepora radians Lamk, 1S36, Ibid., 2 Éd., II, p. 279.
Idmonea radians H. Milne Edw., 1838, "Mem. Crisies", Ann. Sci. Nat. Zool. (2) IX, pp. 214,
217, PI. XII, figs 4 — 4.6 (from one of Lamarck's specimens).
Idmonea radians Busk, 1S56, "Pol. Norway and Finmark", Ann. Mag. Nat. Hist. (2) XVIII,
p. 34 (compared with I. atlantica and distinguished from it).
Idmonea radians Busk, 1875, "Cat. Mar. Pol. Brit. Mus.", III, p. 11, PI. VII, figs 1 — 4
(characteristic figures). (Australia, New Zealand).
Idmonea radians Haswell, 1879, "Cycl. Pol. Port Jackson", Proc. Linn. Soc. N. S. Wales, IV,
p. 350 (Sydney).
Idmonea radians Haswell, 1S80, "Pol. Queensland", Ibid., V, p. 35.
Idmonea radians MacGillivray, 1882, McCOY's "Prodr. Zool. Vict.", Dec. VII, p. 30, PI. LX VIII,
figs 3 — ib (ovicells figured).
Idmonea radians Maplestone, 18S2, "Obs. Living Pol.", Trans. Proc. R. Soc. Vict., XVIII,
p. 51 (colour French grey ; 8 tentacles).
Idmonea radians Waters, 1S84, "Foss. Cycl. Bry. Australia", Quart. J. Geol. Soc, XL, p. 676
(describes lateral windows of ovicells).
Idmonea radians Busk, 1886, "Challenger Rep.", L, p. 10 (Tongatabu, Honolulu).
Idmonea radians Waters, 1SS7, "Bry. N. S. Wales", II, III, Ann. Mag. Nat. Hist. (5) XX,
pp. 203, 255, PI. VI, figs 27, 28 (figures the lateral windows).
Idmonea radians Kirkpatrick, 1890, "Hydr. Pol. Torres Straits", Proc. R. Dublin Soc, (N. S.)
VI, p. 612 (Torres Straits).
Idmonea radians MacGillivray, 1895, "Monogr. Tert. Pol. Vict.", Trans. R. Soc. Vict., IV,
p. 121, PI. XVI, figs 18— i8<r.
Idmonea radians Philipps, 1899, "Polyzoa", WlLLEY's Zool. Res. IV, pp. 441, 449 (Lifu,
Loyalty Islands).
Idmonea radians Neviani, 1900, "Monogr. Gen. Idmonea", Parte I, pp. 26, 41, 46.
Idmonea radians Waters, 1907, " Tubucellaria' ', J. Linn. Soc. Zool., XXX, p. 129 [ovicell
compound, "containing a group of larvae in each of the four to six divisions"].
Idmonea radians Thornely, 191 2, "Mar. Pol. Ind. Ocean", Trans. Linn. Soc. (2) Zool. XV,
p. 156 (Seychelles, Amirante Islands).
Idmonea radians Waters, 1914, "Mar. Fauna Brit. E. Afr.", "Bry. Cycl.", Proc. Zool. Soc,
p. 844, PI. II, figs 6, 7, 8, 10.
" " '| Stat. 37. Sailus Ketjil, Paternoster Islands, o — 27 Metres; coral and coral-sand.
251. J. Stat. 315. Anchorage E. of Sailus Besar, Paternoster Islands, o — 36 Metres; coral and
Lithothamnion.
Also, in the Collection of the Univ. Museum of Zoology, Cambridge : —
Torres Straits (46, 47, 89), A. C. Haddon Coll., Reg. Feb. 24, 1898.
Port Denison, Queensland, Miss E. C. Jelly, Reg. May 24, 1895.
Port Denison, Queensland, from Coll. described by W. A. HASWELL, Australian Mus., Sydney,
Reg. Oct. 23, 1899.
Victoria, Port Phillip Heads, Miss E. C. JELLY, Reg. May 24, 1895.
Lifu, Loyalty Islands, A. WlLLEY Coll., Reg. Mar. 1, 1898.
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140
Zoarium saucer-shaped, attached by a thick short stem, which bifurcates repeatedly to
form a number of narrow branches, of nearly uniform width. The angle between two branches
of a bifurcation is acute. In specimens which have grown vigorously, the two branches formed
by a bifurcation are more or less parallel, and resemble the prongs of a tuning-fork. Branches
subtriquetrous in transverse section, the basal surface slightly convex and the two lateral surfaces
sloping upwards to meet one another in a rounded ridge which runs along the middle of the
frontal surface and unites the peristomes of the median zooecia. In basal view the branches
appear narrow and strap-like, with sharply marked, parallel, straight or slightly sinuous lateral
edges, each of which is formed by the outermost of a series of ridges which occur on this surface
of the branch. These ridges, which are usually about 6 — 8 in number, run longitudinally along
the basal surface. At the growing ends (fig. 8) they appear as longitudinal carinae on the basal
surfaces of the young zooecia. In the interzooecial grooves thus formed are the ordinary zooecial
pores. By the subdivision of the grooves the pores become included in a longitudinal series of
cancelli, each of which contains one or more of the pores. In the older parts of the branches
the openings of the cancelli appear as a single series of moderately large pores, corresponding
with each of the original interzooecial grooves, which are now .separated from one another by
ridges with rounded sides. A similar set of cancelli are developed on the fronto-lateral sides
of the branch and on the ovicells. Series of zooecia oblique, regularly alternating on the two
sides of the branch; 3 or 2 zooecia in each series, the innermost zooecium much the most
prominent, and the outermost one with its orifice not or hardly raised above the level of the
surface of the branch. Ovicells developed just before or at a bifurcation, commonly simple,
but occasionally bifurcating with the branch. The frontal surface of the ovicell is much raised
and is covered with conspicuous cancelli, at the bottom of which are pores of the usual type.
The ovicell corresponds in length with several of the oblique series of zooecia. Between each
two series the wall of the ovicell, on each side, is composed of a flat, thin plate which is
thickly covered with pores; thus forming a series of lateral porous windows, the number of
which corresponds with that of the intervals between the series of zooecia which are involved
in the ovicell. These porous windows develop no cancelli. Each ovicell has a single ooeciostome,
which occurs at either the proximal or the distal margin of one of the lateral windows, and
has the form of a convex hood overarching the window and bearing the slit-like ooeciopore
on its inner side.
This very beautiful species has frequently been described and is characterised by an
unusual number of distinctive features. It seems to be always more or less saucer-shaped, and
to be attached by a central or sub-central stalk. Its branches are arranged in a single tier
(subject to occasional irregularities of growth) ; and from their narrow form and the acute angles
formed by their bifurcations, a large number of branches can be accommodatèd in the space
available. The tuning-fork-like mode of bifurcation (fig. 8) is very characteristic of well grown
colonies. The largest colony observed measures 18 mm. in greatest length; but this measurement
is really a radius of the colony, which may be estimated to have been something like 36 mm.
in diameter when complete. The specimens recorded above from Victoria are much smaller,
the diameter of the largest being only 7 mm., and most of them being not more than 4 or
140
141
5 mm. in that measurement. In all essential respects, including the characteristic ovicells, these
specimens agree completely with those belonging to the 'Siboga' Collection.
The profuse development of cancelli is another very distinctive feature, especially as this
character is not ordinarily found in species of "fdmonea". They are present in all parts of the
zoarium except on the peristomes of the zooecia and on the porous lateral windows of the
ovicells. Both on the basal and on the fronto-lateral surfaces they originate by the subdivision
of the interzooecial grooves. Examination of the arowinsr ends of the branches shows that a
cancellus commonly includes several pores.
The feature which is, however, specially distinctive of the present species is the occurrence
of thin lateral regions in the wall of the ovicell, which have been alluded to above as the lateral
porous windows1). The pores are here much more numerous than in any other part of the
colony ; and they are indeed arranged so closely that there is very little interval between any
tvvo pores. In the 'Siboga' material the ovicell commonly corresponds in length with four of
the oblique series of zooecia on each side (fig. 7). In these cases it follows that there will be
three porous windows (w) on each side of the ovicell; but in ovicells which correspond "with a
smaller number of zooecial series, the number of windows is smaller in proportion. If an ovicell
extends into the bifurcation of a branch, a distal window is generally present, in addition to
the lateral ones. The windows, which are longest in the fronto-basal direction, are usually quite
distinct from one another, as shown in figs 6 and 7 ; and the series of zooecia then extend
along the sides of the ovicell without any obvious modification. In a few cases I have found
that the lateral windows may be confluent along their basal portions ; and the zooecial series
are then interrupted ; a median zooecium occurring in its usual place, the middle zooecium of
the series being suppressed, and the outermost zooecium being found on the basal side of the
confluent part of the windows, separated by a considerable interval from the median zooecium
of the same series.
The lateral windows of the ovicells of this species have already been described by
Waters (1887; see synonymy); and the ooeciostomes have been described by the same author
in a recent paper (19 14). One ooeciostome is present in each ovicell, corresponding with one
of the lateral windows, either on the inner or on the outer side (as determined by the relation
of the ovicell to the preceding bifurcation of the branch). Where three lateral windows are
present on one side I have generally found the ooeciostome on the distal window ; less commonly
on the middle window ; and very rarely 011 the proximal window. The thin perforated plate is
surrounded by a slightly raised margin, and part of the more frontal portion of this margin
is produced into a convex hood (fig. 7, o), which constitutes the ooeciostome. In the majority of
cases this structure is developed from the proximal side of the window, and is directed distally,
as in fig. 7 ; but in a smaller proportion of cases it is developed from the distal side of its
window and looks proximally. The back of the hood of the ooeciostome rests against the
adjacent zooecium, usually the middle one of the series. The ooeciopore is completely concealed
in a frontal view of the branch; but it can be found by arranging the ovicell so as to be able
1) These stvuctures are stated by Waters (1914, p. 845) to be absent in specimens from Zanzibar.
141
I42
to look into the mouth of the ooeciostome, when it is found to be a narrow slit, elongated
transvèrsely. The ooeciostome is thus very similar to that of Tubulipora phalangea, as described
by me in an earlier paper l).
During the development of the ovicell the lateral windows are the parts which calcify
last. The cancellate frontal wall is thus completely formed while the windows are closed merely
by ah uncalcified membrane. I have not obtained specimens in which the complete development
of the normal ovicell can be traced ; but in a specimen from Torres Straits, in which an abortive
attempt to produce an ovicell has been made, two of the median zooecia on one side and three
on the other have each developed a spoon-shaped process which passes towards the middle
line of the branch, at a level considerably raised above that of the general surface of the
branch. The processes have not met one another; but this case may give some indication of the
way in which the roof of the ovicell is normally formed. In this instance one of the branches
immediately beyond the next bifurcation has developed a normal ovicell; but in other cases the
production of an ovicell seems to exhaust the energy of growth, in this respect; no second
ovicell being formed on any branch.
I find comparatively little variation in the present species, but there is some variation
in the form of the orifices. The typical number of zooecia in a series is three ; but two are
not infrequent and four are occasionally observed. The orifices are usually oval, corresponding
with some compression in the peristomes; the longer axis of the oval being in the direction of
the row of orifices of the series. The median peristome is usually long; and those of opposite
sides diverge from one another. The orifice of this zooecium is frequently pointed on its median
side, so that it is not a true oval; and the peristome is often produced into a distinct point
on the same side, in such a way as to resemble a quill-pen. lts outer margin is less often
produced into a point. The next peristome is distinctly shorter than the median one, and in
some cases is produced into a marked point on its outer side, the orifice being then pointed
on that side. The outermost orifice is usually level with the surface of the branch, and no
peristome is present. The middle line of the frontal surface of the branch is generally occupied
by a pronounced thickening band of calcareous matter, from which the median peristomes
diverge. The outer margin of the basal surface of the branch may be slightly serrate,
particularly in young branches.
Me as ure men ts , in a: —
Width of branch, lower part of fig. 8, 500 ;
Width of branch, upper part of fig. 8, 350 ;
Length of ovicell, fig. 6, 1,300;
Longest diameter of proximal lateral window of ovicell, fig. 6, 250 ;
Longest diameter of ooeciostome, fig. 7, 100 ;
Longest diameter of orifices of zooecia, up to 100.
1) Harmer, S. F., 1898, "Dev. Tubulipora", Quart. J. Mier. Sci., XLI, p. 94, PI. VIII, figs. 5, 6.
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143
Tervia Jullien.
Tervia Jullien, 1SS3, uDrag. du Travailleur", "Bry.", Buil. Soc. Zool. France, VII, p. 500
(sep., p. 4).
Idmonea (pars) auctt.
Filisparsa (pars) auctt. (nee D'Orbigny).
The history of the species which have been referred to Tervia is somewhat involved.
The genus was founded by Jullien, in 1883, for four species dredged in the' Atlantic. Two
of these species, T solida and T. discreta have been regarded by Morman j) as forms of T.
irregularis Meneghini. In 1903 Jullien and Calvet'-) admitted that T. folini, one of the
species which had been included in the genus by Jullien in the original account was a synonym
of T. irregularis.
In 1853 D'Orbigny3) founded the genus Filisparsa for several fossil and recent species,
of somewhat varied appearance ; and his name has been accepted by certain later writers, as
by Gregory *) and Pergens 5).
The character by which Tervia was distinguished from Idmonea in the original account
was that some of the zooecia are "isolées, disposées sans ordre sur Ie milieu des branches,
entre les séries latérales". This character by itself might not be considered a sufficiënt reason
for establishing the genus ; but it has been pointed out by Waters 6), in several places, that
the ovicell of Tervia irregularis is "dorsal" in position, while that of Hornera violaeea, var.
tubulosa, or H. tubulosa Busk v), which he refers to Filisparsa, is situated on the front of the
branch. The "dorsal" ovicell of Tervia has been figured by Waters8), Neviani 9) and Jullien
and Calvet 10).
1. Tervia ■ jellyac n. sp. (PI. XI, figs 1 — 3).
Idmonea irregularis Haswell, 18S0, "Pol. Queensland", Proc. Linn. Soc. N. S. Wales, V, p. 35.
Idmonea ? irregularis (pars) Waters, 1S87, "Bry. N. S. Wales", III, Ann. Mag. Nat. Hist. (5)
XX, p. 255 (the specimens from Holborn Island, Queensland).
(n e c Idmonea irregularis Meneghini).
1) NORMAN, A. M., 1909, "Pol. Madeira", J. Linn. Soc, Zool., XXX, p. 2S0.
2) Jullien, J. and Calvet, L., 1903, "Bryozoaires", Rés. Camp. Prince de Monaco, XXIII, p. 114.
3) D'Orbigny, A., 1853, "Pal. Franc Terr. Crét.", V, p. 814.
4) Gregory, J. W., 1899, "Cat. Foss. Bry. Brit. Mus.", "Cret. Bry.", I, p. 66 (admitted provisionally).
5) Pergens, E., 1890, "Rev. Bry. Crét.", Buil. Soc. Beige Géol., III, p. 350.
6) Waters, A. W., 1884, "Foss. Cycl. Bry. Australia", Q. J. Geol. Soc, XL, p. 687; 1S88, "Ovicells Cycl. Bry.", J. Linn.
Soc, Zool., XX, p. 279; 1910, "Mar. Biol. Sud. Red Sea", "Bry. II", Ibid., XXXI, p. 236; 1914, "Mar. Fauna Brit. E. Afr.", "Bry.
Cycl.", Proc. Zool. Soc, p. 843.
7) Busk, G., 1875, "Cat. Mar. Pol. Brit. Mus.", III, p. 19, PI. XVIII, figs 2 — 4. The specimen in question, believed to have
come from the North Atlantic, is preserved in the British Museum (75. 5. 29. 29). The oociostomes, of which two are well shown, have
a considerable resemblance to those of Ttibulipora apcrta\ and the widely open ooeciopores, facing frontally, are also like those of that
species. The zoarium is, however, composed of narrow, bifurcating branches, in which the zooecia are mostly scattered, with but little
indication of the formation of transverse series. It seems to me by no means improbable that the form which I described as Tubulipora
aperta is merely an adnate phase of Hornera tubulosa Busk; although the characters of the ovicell show that the latter cannot be placed
in the genus to which BUSK referred it.
S) Waters, A. W., iSSS, PI. XIV, figs 5, 6.
9) Neviani, A., 1891, "Bri. postplioc. Livorno", Boll. Soc. geol. Ital., X, PI. IV, fig. 21 ; 1905, -Bri. foss. Carrubare", Ibid.,
XXIII, text-fig. 17 on pag. 547.
10) Jullien, J. and Calvet, L.. 1903, PI. XIV, fig. 7.
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'44
? Tervia irregularis Waters, 1914, "Mar. Fauna Brit. E. Afr.", "Bry. Cycl.", Proc. Zool. Soa,
p. $43, PI. IV, fig. S.
285. A.) Stat. 51. Madura Bay, S. point of Molo Strait, 69 — 91 Metres; fine grey sand, coarse
560. A. ) sand with shells and stones.
Also, in the Collection of the University Museum of Zoology, Cambridge : —
Torres Straits, 191. A., A. C. Haddon Coll., Reg. Feb. 24, 1898.
Queensland, Holborn Island, Miss E. C. JELLY, Reg. May 24, 1S95 (now transferred to the
British Museum).
The following specimens, in the Stomafopora-condition, may be noticed in this place.
One or more of them may be the encrusting bases of the present species, but it is probable
that several species are represented, belonging to Entalophora and Tuèuiipora as well as
to Tervia : —
17. T. Stat. 49a. Sapeh Strait, 69 Metres; coral and shells (on Smittia slide, 17. K.).
62. N. Stat. 99. Anchorage off N. Ubian, 16 — -23 Metres; Lithothamnion-bottom.
249. H. Stat. 310. 8° 30' S., H9°7'.5E., Ji Metres; sand with a few pieces of dead coral.
(Cambridge Coll.) Torres Straits, 193, A. C. HADDON Coll., Reg. Feb. 24, 1898.
Zoarium commencing with a " Stomatopora" -Yike. base, giving off erect sterns which are
oval or nearly circular in transverse section. The basal surface of the erect branches is at first
flat, but later becomes covered by a layer of longitudinally disposed kenozooecia which make
this surface convex. Zooecia not arranged regularly in two alternating series, those of opposite
sides beine often continuous across the middle line of the frontal surface of the branch. At the
outer ends of the rows, the zooecia are partly or entirely connate and here resemble the series
of a typical Tubulipora. Ovicell developed on the basal surface of the branch, extending into
a bifurcation, in the angle of which is situated a short ooeciostome which may be visible in
frontal view.
I name this species in honour of Miss E. C. Jelly, from whorri I obtained the ovicell-
bearing specimen here figured, and whose excellent "Synonymie Catalogue of the Recent Marine
Bryozoa" (1889) has been so invaluable an aid to students of recent Polyzoa. It appears to have
been referred to by Haswell and Waters, under the name of Idmonea or Tervia irregularis,
but I think I am justified in separating it from the Mediterranean and Atlantic forms to which
Meneghini's specific name properly belongs. The ovicell of T. jeliyae has much resemblance to
that of an Italian fossil ascribed by Neviani, 1891 x), to Filisparsa variaus Reuss; but I do not
think there is sufficiënt reason for placing the specimens here described under Reuss' species.
The present species is represented only by small fragments. The specimen 285. A. consists
of the base, which is adnate to a small Gasteropod shell inhabited by a Hermit-crab ; and of
two erect sterns of short length. The base by itself might well have been referred to Repto-
tnbigera or Stomatopora -), consisting, as it does, of completely adnate lobes growing over the
shell. These lobes are very irregular in their arrangement, and to some extent confluent with
1) 1. cit.
2) Cf. Waters, A. W., t. cit., p. 844.
144
145
one another. The zooecia here show no uniformity in position. Part of one lobe is formed of two
successive pairs of connate zooecia, which have the relations of those of typical Reptotubigera.
Most of the zooecia of the adnate part are, however, alternate; a meclian zooecium having one
other on each side of it, but at a different level in the length of the lobe. Towards the end
of one of the adnate lobes, the zooecia are assumingf the serial arrangement seen in the erect
sterns. The margins of the lobes may give off kenozooecia, of about the same diameter as the
complete zooecia, which fix the lobe firmly to the shell. Where an erect stem is developed,
some of these kenozooecia1) pass longitudinally up the basal surface of the stem, which, however,
grows more rapidly than the bundie of basal kenozooecia. The branches thus have a flattened
basal surface near their distal ends, but more proximally the kenozooecia grow up the back
of the branch, so as to give this surface a convex shape. The pores are very numerous on
all parts, including the free peristomes. The zooecia of the erect sterns are similar to those of
the Oueensland specimen represented in fig. i ; but many of them have free peristomes, some
of moderate length.
The specimen represented in figs 1,2 is from a Oueensland slide which formerly
belonged to Miss Jelly. It is one of four fragments which consist of erect sterns, of a nearly
cylindrical shape; and there can be no reasonable doubt that they belong to the same species
as the 'Siboga' specimen above described. The branches bifurcate several times, sometimes
at an acute angle, sometimes at an angle of nearly 900. The basal surface is convex and is
marked by distinct longitudinal septal lines, which project as slight ridges, between which the
surface of the zooecia is very gently concave. From the evidence afforded by 285. A. it may
be assumed that this appearance is du e to the kenozooecia which have grown up the back of
the branches from the proximal end of the colony. On the frontal surface the zooecia are
arranged in series, usually with no break in the middle line. In the second series from the
distal end seen, towards the upper end of the figure, in the left hand branch, the transverse
row of zooecia extends in a spiral way across the whole uïclth of the branch, and is completed
by the zooecium projecting to the right ; this zooecium being separated by an interval from
its nearest neighbour. In the series next on its proximal side an interruption in continuity
occurs nearer the middle line of the branch, and in such a position that two of its zooecia
appear to belong to the left side and four to the right side. There is in fact a good deal
of irregularity in the disposition of the series, which have a tendency, however, to extend
across the whole width of the branch, in such a way as to form a curve which is convex
towards the proximal end. At the outer ends of the series, two or more of the zooecia are
commonly connate distally, the peristomes not showing much tendency in this specimen to
become free.
Three of the fragments on this slide possess ovicells, of the type represented in figs 1, 2.
The greater part of the ovicell belongs to the basal surface of the branch (fig. 2), where it
occupies the length of about four zooecial series. At its proximal end the ovicell is no wider
than a zooecium; but it then expands in a pear-shaped way, until it reaches its broadest part,
1) The "canaux de renforcement" of Pergens (1S90).
S1BOGA-EXPEDITIE XXVIII (7. 19
146
which is situated in the emargination of the bifurcation. The basal surface of the ovicell is
only slightly convex; and therefore projects but httle beyond the general level of the basal
surface of the branch. It is covered by very numerous pores; which, as is usual in Cyclostomes,
are more numerous than those of other parts of the colony. In the frontal view (fig. 1) the
distal end of the ovicell appears in the emargination of the branch. lts ooeciostome is a very
short tube, which stands vertically, so that the ooeciopore is easily seen in this view. This
opening may be described as circular, with one part of the circumference flattened. On its
distal side the ooeciostome is in close contact with the peristome of the outermost zooecium
of the series next beyond the bifurcation on the left side. In the other ovicells, the ooeciopore
is concealed by the corresponding peristome and is thus not visible in frontal view, and it is
less circular than in the specimen figured. The ovicell appears to be a modification of one
of the kenozooecia of the basal surface of the branch.
The ovicells of Tervia irregularis have been figured by Waters x). One of the figures
(fig. 5) given by this author, and stated by him to be the normal ovicell of the species, is
elongated, and is not unlike that shown in fig. 2 of this Report, except that it commences
just on the proximal side of a bifurcation and belongs mainly to one of the branches thus
formed. The other (fig. 6) is described as "a short abnormal one"; and is hardly longer than
broad; but it has the same relation to the bifurcation that is shown in my own figure. In
both cases the ooeciostome appears to be visible in basal view. A similar short ovicell is
figured by Jullien and Cal vet, 1903 (t. cit.).
I have compared the forms described in the present Report with specimens of /.
irregularis Menegh. , in the Cambridge and British Museum Collections. Ovicells are ' not
present; but while the general arrangement of the zooecia is similar to that found in the
Oriental and Australian forms, the specimens differ in having their lateral zooecia developed
into strongly projecting wing-like outgrowths of the sides of the branch; each such outgrowth
being formed of several (as many as four or five) zooecia, which are in the main connate, but
have some of their constituent peristomes free at their ends. The pores are less numerous
than in the specimens here referred to the present species. The differences, both of the ovicells
and of the zooecia, seem to be sufficiënt to justify the separation of the forms described in
this Report from the Mediterranean species.
M e a s u r e m e n t s , in y. :
Width of the branch, at the proximal end of fig. 1, 700 ;
Width of the same branch, immediately distally to a row of zooecia which runs transversely
across the entire branch, just before the bifurcation, 1,550;
Diameter of the ooeciopore, 175;
Diameter of the zooecia, 150 — 200 ;
Diameter of the orifices, 150;
Length' of the part of the ovicell shown in fig. 2, 2,550;
Width of the ovicell, at its distal end, 750.
i) Waters, A. W., 1888, t. cit., PI. XIV, figs 5, 6.
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'47
Fam. Horneridae Smitt.
Horneridae Smitt, 1866, "Krit. förteckn. Skand. Hafs-Bry.", II. Öfv. K. Vet.-Ak. Förh., XXIII,
p. 404.
Hornera Lamx.
Hornera Lamouroux, 1S21, uE\-p. Méth.", p. 41.
Hornera Lamouroux, 1827, "Encycl. Méthod.", "Zooph.", p. 460.
1. Hornera spinigera Kirkp. (Fl. XI, figs 8 — 13).
Hornera spinigera Kirkpatrick, 1888, "Pol. Mauritius", Ann. May.. Nat. Hist. (6) I, p. 83,
PI. X, figs i — i/;.
Hornera spinigera Philipps, 1899, "Polyzoa", WlLLEV's ''Zool. Res.", Pt IV, pp. 441, 449 (Lifu).
470. B. Stat. 97. 5°48'.7 N., ii9°49'.6E., 564 Metres; coarse coral-sand.
82. A. Stat. 105. 6°8'N., i2i°i9'E., 275 Metres; coral-bottom.
483. B. Stat. 117. Kwandang Bay, entrance, 80 Metres (chart) ; sand and coral.
112. B. Stat. 156. o°29'.2S., I30°5'.3E., 469 Metres; coarse sand and broken shells.
Also (Univ. Mus. Zool., Cambridge) : —
Lifu, Loyalty Islands, A. WlLLEV Coll., Reg. Mar. 1, 1898 (determined by Miss E. G. PHILIPPS).
Zoarium usually branching in one plane, the branches numerous and close together,
alternate or nearly or quite opposite. Spines developed on many parts, including the peristomes.
Basal surface of the young branches with strong longitudinal ridges, between which the surface
is concave and bears a row of pores. In the older branches the ridges become confiuent,
more numerous and less regular, the pores being still distinguishable. On the frontal surface
the zooecia are arranged in Tuóu///>ora-Yike alternate series, each of which consists of 2 or 3
zooecia, the outermost the most prominent. Peristomes spinous, some of the spines radiating
inwards towards the centre of the orifice. The spines are most developed on the outer zooecia,
and a pair of specially long ones are often produced by the distal margin of the peristome,
and a large spine by the middle of the proximal border. This suboral spine may be greatly
developed, and is sometimes as long as the peristome. Ovicells on the basal surface, extending
along the region of two consecutive branches, their basal surface reticulato-punctate and more
or less strongly carinate. Ooeciostome visible in the frontal view, situated in the axil of a branch.
This very beautiful and characteristic species has been well described by Kirkpatrick
from specimens found at Mauritius; and it has also been recorded from Lifu by Miss Philipps.
It is specially characterised by the serial arrangement of its zooecia and by the spines which
occur on various parts, particularly on the peristomes. In the 'Siboga' specimens the series
are usually bizooecial ; but in that from Lifu three zooecia generally occur in a series. The
outer peristome is long and free, while the inner zooecium has its orifice almost level with
the surface of the branch.
The branches of this species are developed very freely, succeeding one another with but
small intervals between them. They are given oft by both sides of an older branch and may be
alternate, opposite or nearly opposite one another. Most of the branches are in one plane.
At the growing tips (fig. 9) the zooecia are very distinct. Their frontal surface is here
147
148
flat or gently concave, its plane being placed in such a waj' as to project most on the median
side, and falling away towards the outer side of the branch. This flat surface is bordered by
a longitudinal ridge on each side; and these ridges are recognizable as longitudinal ridges
some way down the branch. A few pores occur on the flat surfaces. In the older branches
(fig. 1 2) a considerable amount of thickening is found to have taken place. The middle line
of the branch is the most projecting part; but the ridges have become less regular and more
numerous, and are wider than at first, with evenly roundecl summits. Pores are visible in the
intervals between the ridges, and the external opening of the pore is commonly longitudinally
slit-like. A few sharp spines are developed, here and there, on the frontal surfaces of the
zooecia, principally on the ridges. On the basal surface the spines may be numerous, but are
shorter and conical.
The greatest development of the spines occurs, hovvever, on the peristomes, both of
the inner and of the outer zooecia. They may be regard ed as due partly to the splitting of
the peristome, at its free end, into a certain number of sharp spines, of which two on the
distal border and a stronger, median, suboral spine are usually the largest (fig. 10). The
peristome bears other spines, as well as these, and some of them radiate into the orifice, as
has been pointed out by Kirkpatrick.
The shape of the outer peristomes is variable. In some cases (fig. 9) the peristome
does not widen at its distal end, and the spines are more or less equally developed all round,
although even in these zooecia the suboral spine is generally distinguishable. 112. B. included
two specimens, of very different appearance. In one of these (fig. 1 2) most of the outer
peristomes widen considerably at their distal end, where there is a marked calcareous collar
round most of the orifice, being specially developed proximally and laterally. The suboral
spine is produced from the middle line of this collar, and may be strengthened by a longitudinal
buttress running down the frontal surface of the zooecium. The collar forms a concave shelf-
like surface, starting from the inner aspect of the suboral spine and passing to the sides of
the orifice. In the other specimen (figs 8, 10) from the same dredging the peristomes of
the outer zooecia are greatly compressed ; and they are further remarkable for the enormous
development of the suboral spine, which is compressed and finely denticulate, and may be as
long as the peristome itself. A similar suboral spine may be found on the inner zooecia as well.
In the basal view of young zooecia (fig. 11) a strong longitudinal ridge is generally
discernible along each edge of this surface. Where the two distal spines are large, one of these
ridges is seen to pass into each spine; and the ridges themselves may be plate-like and very
high, so that each lateral border of the basal surface bears a high flange. The basal ridges of
the terminal zooecium of the branch can be traced into the two ridges which run longitudinally
along the back of the young branch (fig. 11). These ridges become joined by others which
belong to the more proximally situated zooecia, and the number is thus increased. In the older
branches the basal ridges are not so regularly longitudinal and they have a more rounded
surface, between them occurring the pores, each of which generally lies in a somewhat lozenge-
shaped depression (upper part of fig. 11), of the kind so frequently seen in species of Hornera.
The outer ends of the pores are generally longitudinally slit-like.
14S
149
Three ovicells are present in the 'Siboga' specimens; one on 112.B. (figs 11, 12) and
the other two on 483. B. They are of the type characteristic of the genus, occurring on the
basal side of a main branch, and belonging to the region of two lateral branches, one on
either side of the main stem (hg. 11). The ovicell is considerably longer than broad and it is
marked by a distinct longitudinal carina which runs obliquely, in the direction of the longest
axis of the organ. From this carina its sides fall off, becoming convex and joining the branch
in such a way that the ovicell is sharply marked off from the neighbouring parts. The whole
surface, with the exception of the carina, is reticulato-punctate ; the pores being situated at the
bottom of fairly deep pits, one of which surrounds each pore. This ovicell is bifid distally,
althoueh this character is not found in the other two. On viewing the branch from the frontal
side (fig. 12), it is seen that the ooeciostome occurs in the proximal axil, and is visible from
this aspect. The tube is not developed on the side towards the axil; but on the opposite side
it is raised and hood-like. The ooeciopore is subtriangular, the apex of the triangle being
directed towards the axil. The ovicells of 483. B. are essentially similar. Both are borne by
one small fragment, and have their ooeciostomes in a corresponding position. The tube of the
ooeciostome is, however, indicated on the axillary side, and the ooeciopore is more rounded.
These ovicells are considerably broader proximally than distally. One of them sends a lobe
along the back of the proximal branch: and its carina, which is much less developed than in
112. B., gives off a branch, nearly at right arigles, extending along this lobe. The other ovicell
shows the same features in a less marked manner.
Measurements, in u. :
Diameter of main branch, fig. n, 700 — 750;
Diameter of left lateral branch, fig. 1 1 , 400 ;
Length of ovicell, fig. 11, 1,900;
Diameter of ovicell, at proximal end, fig. 11, 1,200;
Length of longest peristome, without spines, fig. 10, 425;
Length of longest suboral spine, fig. 8, 525;
Diameter of e.xpanded part of peristome, close to orifice, fig. 12, 150.
Diameter of ooeciopore, fig. 12, 120.
2. Hornera caespitosa Busk. (PI. XI, fig. 13).
Hornera caespitosa Busk, 1875, "Cat. Mar. Pol. Brit. Mus.", III, p. 17, PI. XV, figs 1 — 3.
82. B. Stat. 105. 6°8'N., 121° 19' E., 275 Metres; coral-bottom.
Zoarium with branches which are curved towards the frontal surface, some of them
showing a lateral curvature. Basal surface very convex, the ridges neither regular not prominent,
each bearing a series of granulations. Frontal surface not showing regular longitudinal ridges,
even in young branches; with pores in depressions separated by obscure curved ridges. Zooecia
not arranged in series, their orifices quincuncial. Most of the peristomes are not raised, but
the outer peristomes may be somewhat prolonged and then project at the sides of the branches.
Ovicells wanting.
J49
i50
The present species is represented by a single fragment; obtained, with H. spinigera,
from Station 105. It may be described as a very typical form of its genus, but the material
is hardly sufficiënt to enable it to be determined with certainty. It is probable, however, that
it belongs to IL caespitosa, described by Busk from Cape Capricorn, Oueensland. The ovicells
of a form from near Marseilles referred to H. caespitosa have been described by Calvet *) ;
although it seems not improbable that the specimens figured really belonged to H. frondiculata.
Measurements, in u. :
Diameter of branch, at lovver part of fig. 13, 1,020;
Diameter of branch, at left side of the same, 600 ;
Diameter of orifices, 75 — 100.
Fam. Cytisidae D'Orbigny.
Cytisidae D'Orbigny, 1S54, "Pal. Franc. Terr. Crét.", V, p. 1042.
Cytisidae Pergens, 1890, "Rev. Bry. Crét. figurés par d'Orbignv", Buil. Soc. Beige Géol.,
III, p. 384-
The Family Cytisidae, as defined by Pergens, consists of species in which the colony
consists of free branches or is discoidal, having the orifices placed in groups on one of its
surfaces. The peristomes are not prominent, and accessory pores are present. The ovicells are
rounded swellings placed on the oral surface.
The only species corresponding with this definition which was found among the 'Siboga'
dredgings may be referred to Supercytis D'Orb. : — a type which, according to Pergens, is
assumed by young colonies of forms referable to other genera, in which the branches are not
yet much developed. The occurrence of ovicells in the specimens described below seems to
show, however, that the adult features of the species are more or less developed in a form
which has the Supercy tis-cha.ïa.cters. A number of fossil species of this type are known to
Palaeontologists ; and much difference of opinion exists as to their classification. The material
at my disposal is not sufficiënt to enable me to attempt to reconcile these differences; but for
a critical discussion of the subject I may refer to Gregory's work :) published in 1909.
Supercytis D'Orb.
Supercytis D'Orbigny, 1S54, t. cit., p. 1060.
Supercytis ? Waters, 1884, Toss. Cycl. Bry. Australia", Q. J. Geol. Soc. XL, p. 692.
Supercytis (pars) Busk, 1S86, Challenger Rep., L, p. 28.
Supercytis Waters, 1914, "Mar. Faun. Brit. E. Afr.", "Bry. Cycl.", Proc. Zool. Soc, p. 837.
Fascicidipora (pars) Busk, 1875, "Cat. Mar. Pol. Brit. Mus.", III, p. sy.
As defined by D'Orbigny, this genus consists of pedunculate forms, spreading out above
and giving rise to simple or bifurcate branches diverging horizontally but united at their bases.
1) Calvet, L., 1907, "Bryozoaires", Exp. Sci. Travailleur et Talisman, VIII, p. 47S, PI. XXX, tigs 11, 12.
2) Gregory, J. W., 1909, "Cat. Foss. Bry. Jirit. Mus.". "Cret. Bry.", II, pp. xxiv — xli.
150
i5i
The lower surface is covered by a thin epitheca, while the upper surface bears oblique zooecia.
Ovicells, one or two, more or less inflated, occur in die middle of the upper surface of the colony.
Waters (19 14) points out that 6'. tubigera Busk is not correctly placed in this genus.
1. Supercytis watersi n. sp. (PI. XII, fig. 1).
Supcrcytis': digitata Waters, 1884, t. cit., p. 692, PI. XXXI, figs 22, 26, 27 (fossil, Murray
Cliffs, S. Australia).
Type. 400. A. Stat. 164. i°42'.5 S., i30°4/'.5 E., 32 Metres; sand, small stones and shells.
348. I. Stat. 213. Saleyer Anchorage and surroundings, o — 36 Metres; coral-reefs, mud and
mud with sand.
Zoarium consisting of a nearly cylindrical stalk, expanding into a wide, shallow saucer,
the edges of which are produced into rays. Stalk porous externally, marked by numerous
circular lines of growth. Rays having the same structure as the stalk basally, but on their
frontal surface, corresponding with the concavity of the saucer, honeycomb-like, the smaller
openings being perhaps cancelli. Ovicell central, its roof flat, very porous and distinctly
outlined; traversed by several rows of discrete zooecia, the peristomes of which are short and
disposed vertically. Ooeciostome apparently resembling one of these peristomes.
I name this species in honour of Mr A. W. Waters, who has contributed so much to
our knowledge of recent and fossil Polyzoa, and has apparently described it from an Australian fossil.
The present species is represented by only two specimens, one from each of two shallow
water dredgings. The older specimen is shown in fig. 1, and consists of a massive stalk
expanding distally into a shallow saucer, the edges of which are produced into seven rays.
The base of the colony is not quite complete. The outer surface of the stalk is traversed by
several longitudinal grooves, which separate a corresponding number of rounded elevations
from one another. The surface is further marked by numerous circular lines of growth, some
of which occur in the expanded distal part of the colony. The whole of this surface bears
numerous pores. In the other specimen (348. I.) the longitudinal grooves are absent. This
colony is in a younger state of growth, and longitudinal septal lines are visible; about 5
zooecia being seen in the exposed half. The zooecia which thus form the outer part of the
stalk appear to extend throughout its entire length, each opening by one of the orifices seen
in the cavity of the distal saucer-like part of the colony.
In both specimens, practically the entire frontal surface, with the exception of the central
region, where an ovicell is present, is composed of a honeycomb-like arrangement of calcareous
walls. The larger openings thus delimited are no doubt the orifices of zooecia. The smaller
ones are either the proximal ends of young zooecia or are cancelli, similar to those found in
Lichenopora.
In both colonies the central region is occupied by an ovicell, the roof of which is a
nearly flat calcareous lamina, thickly covered by pores. Through this roof extend the peristomes
of a certain number of zooecia, either singly or, as seen towards the right of the figure, two
together. These isolated peristomes are short but distinct, and stand vertically. In the specimen
ïj1
15^
figured they occur in about four rows, with three or four peristomes in each row. At the
margin of the roof of the ovicell a few peristomes are outlined on their central side, where
they project into the region of the ovicell ; while peripherally they are continuous with the
honeycomb-like arrangement of prismatic spaces which is seen in the rays. Between these
partially isolated zooecia runs a sharply marked, but low calcareous ridge, by which the ovicell
is outlined. In the interval between two rays seen opposite the left half of the upper border
of the ovicell in fig. i , the honeycomb has been closed by the development of a porous
calcareous plate across the opening of each tube. In all other parts of this specimen the
ovicell is in contact, peripherally, with the system of open tubes which form the frontal surface
of the colony, including its rays.
The ooeciostome cannot be distinguished with certainty, but I believe this to be due
to the fact that it has a close resemblance to one of the isolated peristomes of the central
region. The lowest member of the middle vertical row of zooecia, in fig. i, is probably the
ooeciostome.
The ovicell of the younger specimen (348. I.) is similar to the one already described,
but the calcification of its roof is not quite complete. The whole of the rest of the frontal
surface of the colony is honeycomb-like ; and the rays are only just commencing to be indicated.
Measurements, in u. ;
Greatest diameter of the colony, fig. 1, 5,500;
Width of the two broadest lobes, 1,300, 1,100 ;
Width of ovicell, 1,850;
Diameter of orifices, 145.
Supercytis digitata was described by D'Orbigny1) from Cretaceous material. The recent
form to which Busk 2) gave the same specific name (as Fasciculipora digitafa), apparently by
an undesigned coincidence, has usually been regarded as identical with D'Orbigny's species,
although I am not sure that this view is correct. D'Orbigny's fig. 8 represents a colony with
two distinct ovicells. One of these (right of figure) has lost its roof; and, as D'Orbigny
pointed out, it shows "en dedans un canal qui communiqué avec 1'intérieur de la colonie".
This clearly indicates that the ovicell is to be regarded as a modified zooecium ; the aperture
shown by D'Orbigny being the point where the dilated part of the ovicell communicates with
its proximal undilated part. The other ovicell is said to be complete. If this was really the
case, it may be concluded that the ovicell in the Cretaceous form is developed in such a way
as not to include any zooecia. This fact, and perhaps the existence of two ovicells, may be
taken as a sufficiënt reason for distinguishing the recent species from its Cretaceous ally.
The specimen (Brit. Mus. 75. 5. 29. 60) described by Busk in 1875 from Cape Capricorn,
Oueensland, seems to be distinct from the 'Siboga' specimens-, while the figure given by the
same author, in the 'Challenger' Report (1886) of "Supercytis digitata"1 appears to show that
that specimen has an ovicell of a different type, and that it cannot be referred either to S.
1) D'Orbigny, 1854, t. cit., p. 1060, PI. 798, figs 6 — 9.
2) Busk, G., 1875, t. cit., p. 37, PI. XXXIII, lig. i.
152
■53
digitata or to S. watersi. Mr Waters :) has described a Xew Zealand fossil which he considers
"undoubtedly the same as the recent forms found by the 'Challenger' ".
The Japanese Cyclostome described by Ortmann ■) as Hypocytis asteriscus has a close
general resemblance to the present species. The genus was distinguished from Supercytis by
the fact that the zooecia open at the ends of the rays and on their 'Unterseite'. As Ortmann
describes the entire colony as expanding 'oberwarts' into a disc prolonged into rays, I take
this to mean that the orifices are borne on that surface of the rays which I have described as
basal in the 'Siboga' specimens. The surface is said to be "gestreift". Fig. 19^ shows distinctly
outlined zooecia radiating from the centre of the colony, with orifices situated in the peripheral
parts of the rays. It presumably represents a view of the basal surface of the rays; in which
case a small circular area indicated in the middle of the figure probably represents either the
foreshortened stalk or the broken attachment of the stalk to the expanded terminal part of the
colony. If Ortmann's account has been correctly understood, it is hardly possible to regard the
Japanese species as a member of the genus Supercytis.
The Supercytis digitata of MacGillivray and Hall3) is said to have its orifices "on
the lower and not on the upper side"; and in this respect it agrees with Hypocytis.
In its zoarial form, S. watersi has a considerable resemblance to Dcfrancia lucernaria
M. Sars, as shown by the figures given by Busk (1875, PI. XXXIII, fig. 3). It appears to differ
from that species in the characters of the ovicells, those of D. lucernaria being described by
Smitt 4) as swollen vesicles between the zooecial series, elongated in a radial direction.
Fam. Lichenoporidae Smitt.
Lichenoporidac Smitt, 1866, "Krit. fort. Skand. Hafs-Bry.", II, Öfv. K. Vet.-Ak. Förh. XXIII,
p. 405.
Lichenoporidae Pergens, 1890, "Rev. Bry. Ci ét. figurés par d'Orbigny", Buil. Soc. Beige Géol.,
UI, p. 378.
Caveidae (pars) D'Orbigny, 1853, "Pal. Franc. Terr. Crét", V, p. 922.
Galeidae Jullien, 1SS8, "Bryozoaires", Miss. Sci. Cap Horn, VI, Zool., pp. 8, 82.
As defined by Smitt, this Family consists of discoidal Cyclostomata, with a marginal
budding zone and zooecia radiating from a cancellated central reyion. For D'Orbigny's genera
included in this Family, reference may be made to Pergens' work, cited above.
Lichenopora Defrance.
Lichenopora Defrance, 1823, "Dict. Sci. Nat. XXVI, p. 256.
Lichenopora Hincks, 1S80, "Hist. Brit. Mar. Pol.", p. 471.
Disporella Gray, 1848, "List Brit. An. Brit. Mus.", I, p. 138.
Discoporella Busk, 1S59, "Crag Pol.", Palaeont. Soc, p. 115.
Discoporetla Smitt, 1. c.
1) Waters, A. W., 1887, "Tert. Cycl. Bry. New Zealand", o. J. Geol. Soc, XI. III, p. 344.
2) Ortmann, A., 1889, "Japan. Bry.", Arch. f. Naturg., Jahrg. LVI, Bd I, p. 66. PI. II, figs 19.7, 19/».
3) MacGillivray, P. H., 1895, "Monogr. Tevt. Pol. Vict.", Trans. R. Soc. Vict. IV. p. 134, PI. XXII. figs. 1, 2: Appendix,
by T. S. Hall, p. 145.
4) Smitt, F. A., 1S66, "Krit. förteckn." II, Öfv. K. Vet.-Ak. Förh., XXIII, p. 494.
'53
SIEOGA-EXPEDITIE XXVIII a. 20
154
Gregorv ') has criticized the reference of recent species to the genus Lichenopora. He
points out that of the three species originally included in Lichenopora by Defrance, L. turbi-
nata -), an Hocene species, was definitely selected as the genotype by D'Orbigny 3), and that
in this species the orifices are "in elliptical radial bundies, and not in single radial lines".
Recent species which have the latter arrangement are thus wrongly referred to Lichenopora,
and should be placed in Discocavea D'Orbigny; or perhaps (p. 233) in Melobesia Lamx or
Disporella Gray, altered by Busk to Discoporella. Pergens 4), on the other hand, considers that
recent species agree in their general characters with L. furbinata, the genotype of Lichenopora.
I do not feel convinced that the arrangement of the zooecia in radial lines or otherwise
can be adopted conveniently for the generic separation of recent species. These are greatly
in need of further revision ; but the number of species considered in this Report is insufficiënt
for that purpose. It may, however, be pointed out that of the three species here described,
the lïrst has its zooecia in strongly marked radial lines, which have either uniserial or biserial
zooecia; the second has most of its zooecia disposed quincuncially ; while in the third the
zooecia might be described as being arranged in biserial elliptical groups. These facts may
indicate that more than one genus is represented ; but I do not feel able to attempt to arrange
the recent species according to this character. It appears to me probable that a comparative
study of the ovicells might gïve better generic distinctions ; and I may refer in particular to
the great difference, indicated below, between L. verrucaria and L. novae-ze/andiac in the
mode of development of the ovicells. It seems to me best, therefore, to leave the recent
species provisionally in Lichenopora, until some satisfactory method of grouping them can be
shown to exist.
The following notes may be added with regard to the generic names which, according
to Gregory, are available for the recent species : —
(1) Melobesia Lamx, 1S12. — Although, as Gregory points out (p. 233) L. radiata
was referred to Melobesia by Audouin in describing Savigny's figures, it is clear from the
descriptions and figures given by Lamouroux that M. pnstulosa 5) was a Calcareous Alga ;
and indeed Lamouroux gives Corallina as a synonym of J\Lelobesia.
(2) Disporella Gray, 1848°). — The only species included in this sub-genus was
Discopora hispida Fleming, a common British species which is generally included in L^ichcnopora
by writers on recent Polyzoa. It is true that Disporella was given by Gray as a subdivision
of T7ibitlipora\ but- Gregory's criticism (p. 234) that it was not used in a generic or subgeneric
sense does not seem to me convincing. Disporella might thus be available if D. hispida and
its allies should be proved to deserve generic separation.
1) Gregory, J. W., 1909, "Cat. Foss. Bry. Brit. Mus.", "Cret. Bry", II, pp. 247, 233.
2) DEFRANCE, t. cit., p. 257, Atlas, PI. XLVI, figs 4 — 4/1 (reproduced in de Blainville, 1834, '•Man. d'Actinol.", PI. LXVIII,
figs 4 — 4/1. The zoarium of L. turbinata has the form of a funnel, atlached by its stalk-like narrow end, and terminated by a discoidal
snrface hearing the serially arranged orifices and numerous cancelli.
3) D'Orbigny, 1853, t. cit. p. 963.
4) Pergens, E., 1889, t. cit., p. 379.
5) Lamouroux, 1816, "Ili.-t. Pol. Cor. Flex.", p. 315. PI. XII, fig. 2; hettev figured in 1821, '■Exp. Method.", p. 46,
PI. LXXIII, figs 17, 18.
6) Gray, .1. E., 184S, 1. cit.
154
155
(3) Discoporella Busk, 1S591), given by Busk as " Discoporella Gray". — It is not
clear whether this was a deliberate emendation of Gray's name or an accidental misquotation ;
but, as has already been remarked by Jullien -), the form adopted by Busk is not valid,
as D'Orbigny :!) had previously introduced Discoporella for Lunulites umbcllata Defrance (now
usually placed in Cupularia), and two other species. The use of Discoporella in this sense
prevents it from being retained for any member of the Lichenoporidae.
(4) Defrancia Bronn, 1825. — As pointed out by Gregory *), the only species mentioned
by Bronn in the original introduction of the genus is D. (Pelagia) elypcata Lamx. Defrancia
and Pelagia Lamx, 182 1 (preoccupied by Pelagia Pér. and Les.), are thus both synonyms of
Apsendesia Lamx, 1821, if Gregory is right in stating5) that Pelagia elypcata is the young
condition of Apsendesia cristata Lamx, 1821.
(5) Discocavea D'Orbigny, 1853°). — This genus is distinguished by D'Orbigny from
other genera of his Fam. Caveidae by having its zooecia in uniserial radiating lines ; — a
character which does not apply to all recent species usually referred to Lichcnopora.
1. Lichcnopora novae-zelandiae Busk. (PI. XII, figs 6 — 11).
Discoporella novae-zelandiae Busk, 1875, uCat. Mar. Pol. Brit. Mus.", III, p. 32, PI. XXX, fig. 2
(New Zealand).
Discoporella novae-zelandiae Haswell, 1879, "Cycl. Pol. Port Jackson*', Proc. Linn. Soc. N. S.
Wales, IV, p. 353.
Lichcnopora novae-zealandiae Hincks, 1S84, "Contr. Gen. Hist. Mar. Pol.", XII, Ann. Mag.
Nat. Hist. (5) XIII, p. 362 (Mergui Archipelago, Burma; no description).
Lichcnopora novae-zelandiae Ortmarm, 18S9, "Japan. Bry.", Arch. f. Naturg., Jahrg. LVI, Bd I,
p. 65, PI. II, fig. 10.
Lichcnopora novae-zelandiae Thornely, 1905, HERDMAN's "Rep. Pearl Oyster Fish.", XXVI,
"Pol.", publ. by the Roy. Soc, p. 127.
Liclienopora victoriensis Waters, 1889, "Ovicells Lichenoporae", J. Linn. Soc, Zool., XX,
p. 284, PI. XV, fig. 4.
Discoporella holdsworthii Busk, 1875, t. cit., p. 33, PI. XXX, fig. 4.
Lichcnopora holdsworthii Waters, 1889, t. cit., p. 285, PI. XV, figs 7, 8.
Lichcnopora holdsworthii Thornely, 19 12, "Mar. Pol. Ind. Ocean", Trans. Linn. Soc, Zool.,
XV, P. 157.
364. H. Stat. 71. Makassar and surroundings, o — 32 Metres; mud, mud with sand, coral.
1 colony.
568. B. Stat. 105. 6°S'N., 1210 19' E., 275 Metres; coral-bottom. 4 colonies.
108. AB.) Stat. 144. Anchorage N. of Salomakiëe (Damar) Island, 45 Metres; coral-bottom
394. I. ) and Lithothamnion. 8 colonies.
514. A. Stat. 172. Gisser, anchorage between this Island and Ceram-Laut, iS Metres; coral
and Lithothamnion-bottom. 1 colony.
354. A. Stat. 240. Banda anchorage, 9 — 45 Metres; black sand, coral; Lithothamnion-bank in
18 — 36 Metres. 9 colonies.
1) Busk, G., 1859, 1. cit.
2) Jullien, J., iSSS, t. cit., p. 83; cf. also Waters, iSS8, "Ovicells Lichenoporae", J. Linn. Soc, Zool., XX, p. 280.
3) D'Orbigny, A., 1S52, t. cit., p. 472.
4) Gregory, J. W., 1909, t. cit., p. 247.
5) Ibid., 1896, "Cat. Foss. Bry. Brit. Mus.". "Jurassic Bry.", p. 171.
6) D'Orbigny, 1S53, t. cit., pp. 925, 957. The genotype. selected by Gregory, 1909, p. 233. is D. irregn Li'Orb.
'55
156
2o8. C. Stat. 282. Anchorage between Nusa Besi and the N. E. point of Timor, 27 — 54Metres;
sand, coral and Lithothamnion. 1 colony.
202. B. Stat. 282. do, do; reef. 2 colonies.
268.B. Stat. 303 and 60. Haingsisi, Samau Island, O — 36 Metres; Lithothamnion. 1 colony.
239. B. Stat. 310. 8° 30' S., II9°7'.5E., 73 Metres; sand with a few pieces of dead coral.
1 very young colony, growing on a peristome of Entalophora proboscidea (on Entalophora-
slide, 239. A.1).
(?sp.) 251. M. Stat. 315. Anchorage E. of Sailus Besar, Paternoster Islands, o — 36 Metres; coral and
Lithothamnion. 1 colony.
Also (Mus. Zool., Cambridge) : —
Torres Straits, 192, A. C. HADDON Coll., Reg. Feb. 24, 1898. 1 colony.
Ceylon, determined by Miss Thornelv as L. novae-zelandiae, Miss L. R. Thornely, Reg.
Apr. 25, 1906, 72. 1 colony.
Japan, Okinose, off Tokyo, 40 fathoms, A. OWSTON Coll., 5. AN., Reg. June 23, 1902. Many
colonies.
Japan, Tosa, Shikoku Island, T. MlTZOBUCHl, Reg. Sept. 19, 1901. 4 colonies.
New Zealand, determined by Mr T. HlNCKS as L. novae-zealandiae ; HlNCKS Coll., Reg.
May 13, 1899. Several colonies.
Zoarium reaching a diameter of 8.5 mm. in large specimens, but usually smaller. Zooecia
in well marked radial rows, the more median zooecia commonly with much elongated peristomes.
The rays are usually composed of a single series of zooecia, but may be biserial at their
median ends or more rarely throughout their length. Ovicells with a cavity which is continuous
from the first ; sending prolongations into the intervals between the series of zooecia ; with a
porous roof, covered by large cancelli. üoeciostome a short simple tube, with a nearly circular
ooeciopore. Cancelli large, sometimes partially occluded by a cryptocyst-like lamina, and with
numerous pin-like spines radiating into their cavity.
The zoarium varies in form, being sometimes plano-convex, with its marginal lamina
completely adherent to the substratum ; or saucer-shaped, with a free marginal lamina. lts
central region may be depressed (in colonies without ovicells), or this may be the thickest
part. In some cases the colony forms a short cylinder, the free end of which is occupied
mostly by a cancellated region, while the zooecial rays descend the sides of the cylinder. The
number of rays may be at least 40, but new rays are added between the older ones, from
time to time as the growth of the colony proceeds.
The specimens referred to the present species vary greatly in their appearance; as for
instance in the prominence of the peristomes, in the form of the ovicells and in the size of
the openings of the cancelli. The particular form assumed appears to depend largely on the
age of the colony and on the presence or absence of ovicells. I have previously shown l)
that in L, verrucaria the ovicell is usually developed at an extremely early stage in the growth
of the colony, and indeed in many cases from the oldest zooecium but one; that its further
growth is associated with the breaking down of the calcareous septa between the „alveoli" or
cancelli which occur at the bases of the peristomes; and (p. 132) that secondary ovicells may
develop in old colonies. In the species at present under consideration I have no evidence
ij Harmer, S. I.. 1896, "üev. Lichenopora1' ', Quart. J. Mier. Sci., XXXIX, pp. 98 et seq.
156
i57
that ovicells are formed so precociously as in L. verrucaria\ and it seems probable that they
are not as a rule produced until relatively late in the development of the colony, and that in
some cases they do not appear at all. There is here no indication of the extension of the
cavity of the ovicell by the addition of alveoli to its growing margin; and the cavity is frorri
the fïrst a continuous space, as shown in fig. 6. The present species agrees with L. verrucaria,
on the other hand, in its capacity for developing secondary ovicells, at a late stage in its growth.
The variability in the appearance of the colonies appears to depend also on the extent
of the production of ovicells; but it is greatly affected by the varying development of the
cancelli. In colonies which have produced no ovicell, the central region is depressed; while
in fertile colonies this is usually the thickest part.
The extent to which the radiating series of zooecia project is further a variable character.
In some cases, as in fig. 8, the rays are extremely prominent at their central ends, where
they may be biserial, with the peristomes of the two series diverging from one another and
to a considerable extent free. The more marginally situated zooecia diminish rapidly in height
to the outer border of the colony. In other cases, as in fig. ii, the rays project but little;
even the central zooecia being but slightly prolonged, and all the zooecia being arranged in a
sinsfle radial series. The extreme modifications are so different from one another that, in the
absence of other evidence, the specimens might well be referred to two species. But, on the
other hand, colonies from the same dredging may show both these extremes, with other
individuals which are intermediate. I have been unable to make any satis'factory division of
the specimens on the basis of the difference between the projection of the peristomes; and it
appears to be necessary to regard this character as one which is really very variable. The
zooecia are commonly compressed, the peristomes having a corresponding form, and being
often produced into a point on the marginal and on the central side.
The form of the roof of the ovicell is another feature which is subject to variation.
Fig. 9 shows a colony in which the roof has probably been broken. The Moor is thus exposed
and is seen to be composed of a smooth calcareous film which has overspread the underlying
cancelli; a few pores, not represented in the figure, leading through the floor of the ovicell
into the cancelli. The part of the roof which is complete is seen to be composed of two very
convex lobes which are thickly covered by pores. These lobes are seen through the openings
of large cancelli which have been developed superficially to the roof of the ovicell.
Fig. i i shows a very different condition ; the lobes of the ovicell being here roofed by
a nearly fiat calcareous lamina, which is covered by pores. This specimen, taken by itself, might
well be regarded as a distinct species ; but those shown respectively in fig. 6 and fig. 8 are
from the same dredging and agree closely with one another in other characters; while they
show the two conditions of the roof of the ovicell which have just been indicated. The younger
specimen (fig. 6) has two distinct ovicells, of which the one on the lower side of the figure
has at present no roof, but its floor is similar to that of the specimen shown in fig. 9 and
already described. The ovicell of the upper side has a roof which consists of a narrow, flat,
perforated lamina, which is not unlike that of fig. 11. In fig. 8, from the same Station,
the growth of the ovicell is nearly complete; but its roof, seen through the widely open
i57
158
cancelli, consists of broad, convex, porous lobes which are similar to those of fig. 9. Fig. 8 is
a colony in which the central zooecia of the rays are specially elongated and have a very
distinctly biserial arrangement. But in fig. 10, representing a specimen which agrees with fig. 1 1
in its uniserial, slightly prominent zooecial rays, the lobes of the ovicell are not flat-roofed,
as in that specimen, but are just as convex as in fig. 9.
It does not seem easy to give any other explanation of these differences than the
assumption that the form of the ovicell is variable. It appears probable, however, that the
ovicell has to adapt its form to the space which is available. Thus the colony shown in fig. 1 1
has a very small projection of its zooecial rays ; while it is growing on the convex surface of
a cylindrical Coral-branch of less diameter than itself. Owing to the latter reason, the basal
surface of the colony is very concave, the edges of the Lichcnopora being reflected round the
Coral. This position would naturally result in some stretching of the central region of the colony,
and would thus tend to prevent it from increasing greatly in thickness. The slight projection
of the zooecial rays would operate in the same direction by not allowing much vertical room
for the growth of the ovicell ; and it is perhaps owing to the combination of these two conditions
that the roof of the ovicell is unusually flat. In fig. 8 the conditions are very different. Although
this colony too is growing on a cylindrical object of less diameter than its own, its basal lamina
is for the most part free; being only attached along a narrow line to its substratum. There is
thus no interference with the growth of the colony in vertical height. As the peristomes are,
moreover, unusually long, there is plenty of room for the ovicell to attain a greater vertical
height than in the other case ; and the roof is accordingly very convex, with a correspondingly
large perforated surface.
In some cases there is evidence of the development of what I have previously termed
secondary ovicells ; that is, ovicells which are formed at a late stage in the growth of the colony.
Thus on slide 108. AB., one of the colonies has a normal ovicell in its central region; and, in
addition, a much smaller ovicell, with an ooeciostome which has developed, in a more superficial
position, between the outer ends of two or three of the zooecial rays. The ovicell shown in
fig. 9 may also be regarded as a secondary one, since it is being developed in a colony which
has already reached a considerable vertical thickness.
The ooeciostome varies little in form, although its position is not constant. In fig. 8 two
ooeciostomes (o) are seen, at two points on the margin of the ovicells, of which it may be assumed
that at least two are present. Each is a short, simple tube, terminated by a nearly circular
ooeciopore. It is usually circular in section, but is occasionally compressed. In most cases the
ooeciostome is found on the central side of one of the zooecial rays, or between the central
ends of two of them. The number of ovicells is apparently always small (not more than two or
three); but the development of cancelli generally prevents the line of junction of two contiguous
ovicells from being visible; and the number of ovicells has to be inferred from that of the
ooeciostomes.
Certain colonies are found to have reached a large size without showing any tracé of
an ovicell. A specimen of this kind, on slide 354. A., has a diameter of 3.2 mm. Its central
region is depressed, and the entire surface which is not occupied by zooecial rays is covered
15S
159
by large, open cancelli through which no tracé of ovicell can be seen. It is not impossible that
colonies of this type have developed male sexual organs only, though I have no evidence on
this subject. The majority of the colonies are, however, fertile.
The marginal lamina is generally broad : — an indication of the fact that growth is not
complete. In some instances, as in certain specimens growing on shells, it is closely adnate to
its substratum. An incidental advantage of this arrangement is well seen on slide 568. B.1, where
a colony is growing over an area occupied by an encrusting Cheilostome. lts marginal lamina
has overspread the Cheilostome, occluding its orifices, and thus removing the competition of
what might have been an inconveniently near neighbour. In one or two cases, however, the
marginal lamina and some of the peripheral zooecia have been covered by an encrusting
calcareous Alga.
In many other instances the marginal lamina has become completely free from the
substratum ; and it may then grow into the saucer-shaped edge which is also found in other
species of the genus.
The cancelli are occupied; — or at least some of them in each colony; - - by the well
knovvn radiating, pin-like spines, as shown in fig. 7. Examined with a sufficiently high power, the
head of the pin is found to be minutely denticulated, or aster-like. In many cases the cancelli
remain widely open ; but in others a cryptocyst-like calcareous lamina develops in an iris-like
way at a level slightly lower than the edges of the cancellus. The orifice of the cancellus may
be reduced in this manner to a very small opening; and colonies in which this process has
taken place may at first sight look very different from those with open cancelli. But even in
these cases the cancelli may be seen to have the normal size, since their original edges still
project as slight ridges above the level of the iris-like cryptocyst. In some of the specimens
from Japan (5. AN) the ridges separating the cancelli are specially high and may be produced
into spinous projections at their nodal points (where the walls of several adjacent cancelli unite).
In some of these specimens the marginal side of the orifices may be produced into a sharp
point ; while in some of those from New Zealand the central side of the orifices may be prolonged
into one or two points, in addition to that on the marginal side. These specimens, from Japan
and New Zealand, are so similar to the 'Siboga' specimens in the characters of the ovicells
that their reference to the same species seems fairly certain.
Some of the 'Siboga' specimens show distinct indications of regeneration of the entire
colony : - - a process which is known to occur in other species of the genus 1). This is specially
well seen in some large colonies (as much as 8.5 mm. in diameter) mounted on slide 568. B\
In one or two of these colonies, a smaller disc has developed from a part of a larger zoarium ;
and the regenerated clisc is produced from one side of the older part of the colony in such a
way that it is not in the least concentric with it, but leaves part of the old surface exposed,
while the remainder is covered by the regenerated, excentric disc.
The specimen 251. M. is very doubtfully referred to the present species. Its rays are rather
more biserial than usual, and more nearly approach the centre of the colony. The characters
1) Cf. Pergens, E., 1890, "Rev. Bry. Crét. figurés par D'Orbigny", Buil. Soc. Beige Géol. III, p. 319.
IS9
ióo
of the ovicell, ooeciostome and cancelli are, however, much as in other specimens which are
believed to belong to the present species.
M easuremen ts , in a: —
Greatest diameter of the largest colony, 56S. B.3, 8,500;
Greatest diameter of the colony, fig. 11, 4,600;
Greatest diameter of the colony, fig. 10, 2,900;
Greatest diameter of the colony, fig. 9, 3,200;
Length of longest peristome, fig. 8, 1,000 ;
Greatest diameter of ovicell, fig: 1 1 , 1,850;
Length of longest zooecial series, fig. 11, 1,200;
Width of zooecial series, fig. 11, 80 ;
Average diameter of openings of larger cancelli, fig. 10, 130;
Diameter of ooeciopore, fig. 8, 75.
The determination of the species of Lichenopora is admittedly very difficult ; and it is
probable that the present species has been described under several specific names. Forms like
the colony shown in fig. 8 have a considerable resemblance to L. radiata, originally described
by Audouin l) as Melobesia radiata, on the basis of Savigny's excellent figures -). In these
figures the rays are represented as being uniserial, with the central zooecia greatly elevated,
and longer than those which occur nearer the margin. The existence of an ovicell or ovicells
is clearly indicated in Savigny's fig. 3*; but it appears from the figures that the cancelli are
smaller than in L. novae-zelandiae, from which it is probably distinct.
Mr Waters 3) has figured the ovicell of a form referred by him to L. novae-zelandiae,
which differs greatly from the ovicells described in the present Report. A central smooth region,
not covered by cancelli, is represented as being surrounded by a sharply marked calcareous
ridge. This has some resemblance to the ovicells of specimens from Ceylon described by Busk
as Discoporella holdsworthii (see below).
L. novae-zelandiae has been recorded from the Azores by Jullien and Calvet *) but in
the absence of figures it is perhaps permissible to doubt the correctness of the reference.
In his paper on the ovicells of Lichenoporae, Waters 5) describes an ovicell which
corresponds closely with those found in the lSiboga' specimens; but he refers the specimens
to L. victoriensis, a new name for L. reticulata MacGillivray, preoccupied. It appears to me
probable that this is really L. novae-zelandiae.
The ovicell figured by Waters g), in the same paper, as that of L. echinata MacG.
develops in the same manner as that represented in my own fig. 6; but MacGillivray's species,
1) Audouin, V., "Explication", "Pol.", 1826, p. 235; Ed. 2, 182S, p. 60. The specimen (Biït. Mus. 90.3.24.54) from Torres
Straits referred by Mr Kirki-atrick (1S902, p. 612) to Lichenopora radiata has strongly marked radial ridges composed of 2 — 3 series of
connate zooecia, and has much resemblance to Radiopora cristata Busk (1S75, p. 35).
2) SAVIGNY, J. C, «Descr. Egypte", PI. VI, figs 3"— 3*.
3) Waters, A. W., 1887, «Bry. N. S. Wales", III, Ann. Mag. Nat. Hist. (5) XX, p. 261, PI. VII, fig. 8.
4) Jullien, J., and Calvet, L., 1903, "Bry.", Rés. Camp. Prince de Monaco, XXIII. p. 164.
5) Waters, A. W., 1889, t. cit., p. 2S4, PI. XV, fig. 4.
6) Ibid., 1889, t. cit., p. 282. PI. XV, fig. 6.
160
iói
which bears numerous fine spines, seems to be distinct from L. novae-zelandiae. The specimen
referred by Waters to L. californica l) has, on the other hand, an ovicell which closely
resembles that shown in my own fig. 1 1 . Whether Mr Waters' specimen really belongs to
L. californica ma)- be regarded as doubtful. It agrees with the form so named by Busk 2) in
having biserial or even triserial rays ; but it should be noted that the Unicavca ca/i/oruica of
D'Orbignv was described as a member of a genus characterised by having uniserial rays. The
form recorded by Miss Robertson 3) under the same name has also pluriserial rays, but it does
not seem to correspond in the characters of its ovicell with the specimen figured by Waters.
The uniserial arrangement of the zooecia in the 'Siboga' colony is a reason for not referring
the specimen shown in fig. 1 1 to the species described by Busk, Waters and Miss Robertson
as L. californica. L. holdsworthii Busk, described from Ceylon, has, on the other hand, a
close resemblance to some of the 'Siboga' specimens referred by me to L. novae-zelandiae. In
the original description *) of " Discoporella holdsiuorthii', Busk makes no reference to ovicells,
although these structures are present in two of the colonies from Ceylon on a slide (Brit. Mus.,
75. 5. 29. 48*) belonging to the collection on which his Catalogue was based. In one of these
colonies the ovicell occupies a central, much depressed region, surrounded by a high circular ridge
formed by zooecia and cancelli. lts porous roof is flat and has a close general resemblance to
that shown in my own fig. 1 1 . Five main lobes, two of which divide into secondary lobes
by a single bifurcation, are separated by central prolongations of the cancellated region. An
ooeciostome, with a large, nearly circular ooeciopore, occurs among the cancelli just outside
the flattened porous roof. In the raised circular ridge surrounding the central depression occurs
a secondary ovicell of the same general type. The other fertile colony has a cylindrico-truncate
form, terminated by a flat region containing one or perhaps two ovicell-roofs of the same
general character as in the other colony.
Taken by themselves Busk's specimens would suggest that L. holdsworthii is distinct
from L. novae-zelandiae; but the study of the material considered in this Report indicates that
this is probably not the case, and that the differences between the two types are due to variation
in the extent to which the cancelli cover the ovicell. It may be remarked that pin-spines are
well developed in Busk's specimens.
2. Lichenopora buski n. sp. (PI. XII, figs 4, 5).
Discoporella ciliata Busk, 1S75, uCat. Mar. Pol. Brit. Mus.", III, p. 31, PI. XXX, fig. 6;
PI. XXXIII, fig. 4.
(nee Discopora ciliata Busk, 1855, "Zoophytology", Q. J. Mier. Sci., III, p. 256.)
Discoporella ciliata Haswell, 1879, "Cycl. Pol. Port Jackson", Proc. Linn. Soc. N. S. Wales,
IV, p. 354-
Lichenopora ciliata Waters, 1887, uBry. N. S. Wales", III, Ann. Mag. Nat. Hist. (5) XX,
p. 263, PI. VII, fig. 5.
1) Waters, A. W., 1889, t. cit., p. 282, PI. XV, fig. 1.
2) Busk, G., 1875, t. cit., p. 32, PI. XXX, fig. 5.
3) Robertson, A., 1910, "Cycl. Bry. W. coast N. America", Univ. Calif. Publ. Zool., VI, p. 261, PI. XXV, fig. 48.
4) Busk, G., 1875, t. cit., p. 33, PI. XXX, fig. 4.
161
S1BOGA-EXPEDITIE XXVIII a. 21
IÓ2
Lichenopora ciliata Waters, 1889, "Ovicells Lichenoporae", J. Linn. Soc. Zool., XX, p. 283.
Lichenopora ciliata, L. hispida and L. verrucaria Philipps, 1S99, Wille Y's Zool. Res., Pt IV,
p. 441.
394. P. Stat. 144. Anchorage N. of Salomakiëe (Damar) Island, 45 Metres ; coral-bottom and
Lithothamnion. 1 colony.
(?sp.) 568. E. Stat. 105. 6°8'N., I2I°I9'E., 275 Metres; coral-bottom. 1 colony.
Also (Univ. Mus. Zool., Cambridge): —
Torres Straits, A. C. Haddon Coll., 6, Reg. Feb. 24, 1898. 1 colony.
Torres Straits, A. C. Haddon Coll., 81, Reg. Feb. 24, 1898. 1 colony.
Japan, Uraga Channel, 30 fathoms, A. Owston Coll., 7. Q., Reg. June 23, 1902. 3 colonies.
Japan, Tosa, Shikoku Island, T. MlTZOBUCHl, Reg. Feb. 1, 1901. 1 colony.
Japan, from rooting spicules of Euplectella marshalli, Prof. K. MlTSUKURI, Reg. Sept. 23,
1896. 1 colony.
Lifu, Loyalty Islands, A. Willey Coll., Reg. Mar. 1, 1898. Determined by Miss Philipps as
L. ciliata. 1 colony.
Lifu, Loyalty Islands, A. Willey Coll., Reg. Mar. 1, 1898. Determined by Miss Philipps as
L. hispida. 1 colony.
Lifu, Loyalty Islands, A. Willey Coll., Reg. Mar. 1, 1898. Determined by Miss PHILIPPS as
L. verrucaria. Two slides, 1 colony on each.
Zoarium reaching a diameter of 3.9 mm. Zooecia quincuncially arranged in the marginal
part of the colony, sometimes in uniserial rays centrally; the orifices of those situated in the
rays sometimes separated by one or two small cancelli. Peristomes produced on their central
sides, pointed like the nib of a pen, or truncate, or fringed into several fine spines. Small
pointed spines may also occur on the outer sides of the peristomes or, more rarely, on the
septa between the cancelli. Ovicells usually small, their roof porous and covered with irregular
ridges, but not by typical cancelli. Ooeciostome a simple tube, terminating in a large, nearly
circular ooeciopore. Cancelli not usually separated by narrow, angular septa, their openings
variable in size and often much reduced by the development of a cryptocyst; pin-like spines absent.
Two colonies from the 'Siboga' Collection are included in the list given above. Of
these, 568. E. is in poor condition and is very doubtfully referred to the present species. The
remaining specimens in the list form a fairly uniform series, in which, however, a certain
amount of variation is shown, particularly in the form of the peristomes and in their tendency
to become united into rays near the central part of the colony.
The present species seems to be characterised by having its more marginal zooecia
isolated and arranged quincuncially, without any attempt to form radial rows. In the more
central part of the colony a few zooecia may be united to form definite uniserial rays, although
successive peristomes are usually separated from one another by a small cancellus or more
commonly by a pair of these structures. This condition is shown by both the 'Siboga'
specimens. In 394. P. (fig. 4) the central border of the peristome is usually elevated into a
single point, resembling the nib of a pen, though some of them have the corresponding part
of the peristome truncated, with a tendency to develop a short spine from each angle of the
truncated end. From this raised central part, the lateral edges of the peristomes slope down
to the general level of the surface of the colony, so that the marginal border is not at all
raised. A few sharp spines, of no great length, are developed from various parts of the outer
162
163
wall of the peristomes. The marginal cancelli are fairly large, and are at first separated from
one another by narrow septa with sharp edges. In the more central regions the cancelli cease
to be separated by angular septa, and their openings are reduced by the development of a
concave iris-like cryptocyst. No ovicell seems to be present in this specimen.
Of the specimens from Torres Straits, N" 6 has nearly all its zooecia isolated and
quincuncial, only one or two near the centre showing some disposition to assume a radial
arrangement. Their peristomes are greatly elevated centrally, but fall off sharply laterally, so
that the orifices have practically no lateral or distal raised wall. The central raised part of
the peristome is either truncated, the edge forming a segment of a circle without spines; or
its edge is fringed out into from three to five long delicate spines. The cancelli have their
openings much reduced throughout the colony. Spinules are borne by the outer walls of the
peristomes, as in 394. P. The marginal lamina is in this case free and upturned at its edge,
so as to form a saucer; and, as has been noticed in some of the other specimens, its surface
has a finely granular texture. In the other Torres Straits specimen (81) it is for the most part
closely adherent to its substratum.
The specimen from Tosa, Japan, has a close resemblance, in most respects, to N° 6
from Torres Straits. The internal edge of the cryptocyst of the cancelli may be very finely
denticulate; a feature which is barely indicated in 394. P. ('Siboga'), but is well marked in
several of the other specimens. None of the colonies show any of the pin-like spines which
are so conspicuous a feature of L. novae-zelandiae. The Tosa colony shows a well marked
ovicell (fig. 5), which has some resemblance, in form, to that of certain species of Bercnicea,
being transversely elongated and giving off an ooeciostome [o) from the middle of its marginal
border. The ooeciostome is a simple tube, which at first runs horizontally, but soon curves,
so that its simple, nearly circular ooeciopore faces directly frontally. The ovicell, which exhihits
no tendency to run into lobes between the zooecia, is more or less outlined by a series of
ordinary cancelli, but its roof is not covered by typical cancelli. This part of the ovicell is,
however, traversed by irregular, low ridges, between which are a moderate number of pores,
each pore being as a rule sunk at the bottom of a slight depression. The roof of the ovicell
is more convex than the adjacent part of the zoarium. The remainder of the central part of
the colony is occupied by a somewhat similar convex surface, which is, however, simply
granular and does not seem to be porous. I cannot distinguish an ooeciostome in connexion
with this part, and I am unable to decide whether it represents a second ovicell or not. There
is nothing to show whether its cavity is continuous with the ovicell, which has a porous roof;
but it is not impossible that it my be merely a part of that ovicell.
Of the three specimens on the Japanese slide J.Q, one has no ovicell; but in each of
the others the central part of the colony is occupied by an ovicell of the type already described,
except that it sends out short lobes, peripherally, between the more centrally placed zooecia.
In one of these colonies the ooeciostome is very distinct ; and in the same colony the ridges
on the roof of the ovicell are produced, here and there, into spinous projections. The cancelli
in these specimens have their openings hardly reduced by cryptocyst. Some of the zooecia are
arranged in uniserial rays, the zooecia of which may be contiguous.
163
164
The remaining Japanese specimen (Prof. Mitsukuri) has the openings of its cancelli
greatly reduced by cryptocyst.
Several of the specimens from Dr Willey's Lifu Collection seem to be referable to
the present species. The colony determined by Miss Philipps as L. ciliata has some of its
zooecia arranged in well marked uniserial rays, although the more peripheral zooecia are, as
usual, quincuncial. Many of these are fringed out into spines at their free ends; and an ovicell,
with an ooeciostome, both of the type found in the Japanese specimens, are present.
A colony, from the same Collection, referred by Miss Philipps to L. hispida, and two
colonies referred by her to L. verrucaria seem to belong to the same species.
M e a s u r e m e n t s , in \j.\
Greatest diameter of colony, fig. 5, 3,900;
Greatest diameter of colony, fig. 4, 3,500;
Diameter of peristomes, fig. 5, 100;
Diameter of openings of cancelli, fig. 4, 50 — 60.
The species here considered appears to be identical with Discoporella ciliata Busk, 1875,
(Cape of Good Hope, New Zealand) ; and with the form described under the same specific
name by other authors. It does not seem to have been noticed, even by Busk himself, that
the specific name is pre-occupied by Discopora ciliata Busk, 1855 (see synonymy), a Northerrt
form clearly referable to Lichenopora and perhaps a form of L. verrucaria. It seems necessary,
therefore, to introducé a new designation, and I have accordingly named it in honour of Busk;
the type-specimen being the slide from the Cape of Good Hope preserved in the British Museum
under the Register-number 99. 7. 1. 519. Mr Waters (1887) has indeed suggested that L. ciliata
Busk, (1875) may be identical with L. verrucaria; but the account which he gives of its ovicell
shows clearly, I think, that this view cannot be maintained; and in any case the ovicell of the
specimens described in the present Report is widely different from that of L. verrucaria.
3. Liclienopora ? mediterranea Mich. (PI. XII, figs 2, 3).
Lichenopora mediterranea Michelin, 1840—1847, "Iconogr. Zoophytol.", p. 68, PI. XIV, figs $a, $è.
Discoporella mediterranea Busk, 1875, "Cat. Mar. Pol. Brit. Mus.", III, p. 33, PI. XXXIV, fig. 4.
(nee Liclienopora mediterranea (nomen nudum), Blainville, 1834, "Man. d'Actinol." p. 407.)
? Discoporella porosa Haswell, 1879, "Cycl. Pol. Port Jackson", Proc. Linn. Soc. N. S. Wales,
IV, p. 354-
? Liclienopora wanganuiensis Waters, 1887, "Tert. Cycl. Bry. New Zealand", Q. J. Geol. Soc.,
XLIII, p. 346 (with text-fig.).
565. A. Stat. 125. Anchorage off Sawan, Siau Islands, 27 Metres; stone and some Litho-
thamnion. 1 colony.
568. C. Stat. 105. 6°8'N., 1210 19' E., 275 Metres; coral-bottom. 2 colonies.
? sp.) 56S. D. Do. 1 colony.
Of the specimens above indicated, 565. A. is a colony in poor condition. It is represented
in fig. 3 ; and, while agreeing with L. novae-zelandiac in the serial arrangement of its zooecia,
it differs from that species in having the rays biserial throughout, and in the form of its
164
i65
cancelli. The rays do not project much and the zooecia are completely connate. It is quite
possible, however, that their length was really greater than is indicated by the present condition
of the specimen and that their distal parts have been worn off. The rays are disposed in
much the same manner as the septa of a Madreporarian Coral ; larger primary rays alternating
with smaller secondary rays. The cancelli are conspicuous and have rounded openings, separated
from one another by thick septa which are not angular at their edges as in L. novae-zelandiac.
The nodal points of these septa have a tendency to be raised into short rounded tubercles.
Pin-like spines project into the cancelli (fig. 2), as in the othér species. Ovicells cannot be
distinguished.
The slide 56S. C. contains two worn colonies, the larger of which has a marked biserial
arrangement of its very prominent zooecial rays l), which are, however, more massive than in
565. A. The cancelli resemble those of that specimen. The other colony is probably a younger
condition of the same species. One of the rays is biserial and massive ; another is biserial but
narrower; and the others are uniserial.
568. D. is a young colony, in better condition, but it has a uniserial arrangement of
its zooecia. The cancelli are like those of the other specimens but their cavity is constricted
by an iris-like cryptocyst, lying at a deep level. In this specimen some of the septa between
the cancelli are prolonged up the zooecia as longitudinal ridges.
Measureraents, in u. :
Greatest diameter of colony, fig. 3, 3,000;
Length of longest zooecial series, fig. 3, 9,50;
Width of zooecial series, fig. 3, 150 — 180;
Average diameter of openings of larger cancelli, fig. 2, 100.
Busk gives this species as Discoporella mediterranea Blainville; but a reference to the
"Manuel d'Actinologie-' shows that de Blainville's name was a nomen nudum, asno diagnosis
of any kind is given, though the species is said to occur, living, in the Mediterranean. The
generic characters as formulated by de Blainville included the statement that the "cellules"
are irregularly scattered; which is by no means in accordance with the strongly marked serial
arrangement shown by Busk. De Blainville even expresses the conviction that the Lichenopora
of Defrance are merely the young stages of a form like " Retcpora reticulata' , which, as
shown by his synonymy (p. 406), is Frondipora verrucosa. As the name given by this author
is a nomen nudum, it makes no difference what was intended ; and Michelin's name may
be considered valid. It is not clear whether Michelin's figures are taken from a recent or a
fossil specimen.
I refer the 'Siboga' specimens with considerable hesitation to Michelin's species, which
seems to be the same as that figured by Busk.
1) As in L. imptrialis Orlmann, 1S89 ("Japan. Bry.", Arch. f. Natuig., Jahig. LVI, Bd I, p. 64, PI. IV, fig. 25), although in
this species the rays appear to be uniserial.
l65
166
Distribution of Cyclostomata.
STATION
DEPÏH in Metres
SPECIES
37
49a
5i
53
59
60 & 303
64
77
80
81
97
99
105
o — 27
69
69—91
0—36
390
0—36
0—32
0—32
59
40—50
34
564
16—23
275
116
72
117
80
119
1901
125
27
129
23—31
133
0—36
139
397
144
45
Crisina radians.
(Undetermined, in Stomatopora-condition).
Tervia jellyae.
Tubulipora atlantica.
Entalopliora proboscidea (?).
Tubulipora pulcherrinta.
Crisia elongata.
Tubulipora pulcherrinta.
Liclienopora novae-zelandiae.
Entalopliora proboscidea.
Liclienopora novae-zelandiae.
Crisia elongata.
Crisia kerguelensis.
Tubulipora atlantica.
Tubulipora pulcherrinta.
Crisia elongata.
Entalopliora delicatula (?).
Tubulipora concinna.
Tubulipora atlantica .
Tubulipora pulcherrinta.
Crisia elongata.
Tubulipora atlantica (?).
Hornera spinigera.
Tubulipora pulcherrinta (?).
(Undetermined, in Stowatopora-condition).
Crisia elongata.
Berenicea sarniensis.
Tubulipora atlantica, var. jlexuosa.
Tubulipora pulcherrinta.
Hornera spinigera.
Hornera caespitosa.
Liclienopora novae-zelandiae.
Liclienopora buski (?).
Liclienopora mediterranea (?).
Tubulipora pulcherrinta.
Tubulipora pulcherrinta.
Hornera spinigera.
Tubulipora cassiforniis.
Liclienopora mediterranea (?).
Tubulipora pulcherrinta.
Crisia elongata.
Berenicea lineata.
Crisia elongata.
Entalopliora intricaria (?).
Tubulipora atlantica.
Tubulipora pulcherrinta.
Liclienopora novae-zelandiae .
Liclienopora buski.
166
i6-
STATION
DEPTH IN Metkes
SPECIES
I56
164
172
204
213
240
248
250
251
257
273
274
282
301
305
310
315
469
32
75—94
0—36
9—45
o— 54
20—45
204
o — 52
13
57
27—54
22
113
73
-36
321
82
Tubitlipora atlantica, var. flexuosa.
Tubulipora cassiformis.
Hornera spinigcra.
Crisia clongata.
Crisia cuneata.
Bntalophora intricaria (?).
Tubulipora pulcherrima.
Supercytis watersi.
Lichenopora novae-zelandiae.
Berenicea sarnicnsis.
Crisia elongata.
Entalophora delicatula.
Supercytis watersi.
Crisia elongata.
Entalophora intricaria (?).
Berenicea lineata.
Tubulipora concinna.
Tubulipora atlantica.
Tubulipora pulcherrima.
Lichenopora novae-zelandiae.
Tubulipora atlantica.
Tubulipora pulcherrima.
Tubulipora pulcherrima.
Tubulipora pulcherrima.
Crisia elongata.
Tubulipora pulcherrima.
Crisia elongata.
Entalophora delicatula.
Tubulipora atlantica.
Crisia elongata.
Crisia elongata.
Entalophora intricaria (?).
Tubulipora pulcherrima.
Lichenopora novae-zelandiae.
Entalophora delicatula.
Tubulipora pulcherrima.
Crisia elongata.
Entalophora proboscidea.
Berenicea sarniensis.
Reptotubigera philippsae.
Tubulipora pulcherrima.
Lichenopora novae-zelandiae.
(Undetermined, in Stoma/opora-conè\\.\on)
Crisia elongata.
Entalophora proboscidea.
Entalophora intricaria (?).
Tubulipora atlantica (?).
Crisina radians.
Lichenopora novae-zelandiae (?).
Crisia kersruelensis.
167
i68
The preceding list gives a complete statement of all the specimens of Cyclostomata
which I have found among the 'Siboga' dredgings. It will be seen that members of this group
have been discovered in 40 Stations out of about 135 Stations at which Polyzoa were obtained;
or almost the same as the number (42) at which either Entoprocta or Ctenostomata, or both
these groups together, were found (cf. pp. 92 — 94). It exceeds the total for either group separately;
Entoprocta having been found at 30 Stations, and Ctenostomata at 29. Cyclostomata were
recorded in company with Entoprocta at 2 1 Stations ; and with Ctenostomata at 1 6 Stations.
Of these sixteen, none yielded less than 13 species of Polyzoa (mostly Cheilostomata); while
at each of the other ten Stations more than 30 species of Polyzoa were discovered.
These facts are further elucidated by the following table ; in which are set out all the
Stations at which Cyclostomata were found in company with Entoprocta or with Ctenostomata,
Stations at which Cyclostomata were found with Entoprocta and Ctenostomata.
STATION.
DEPTH ,
IN METRES.
ENTOPROCTA.
CTENOSTOMATA. CYCLOSTOMATA.
Species of POLYZOA
in geneval (including
CHEILOSTOMATA).
53
0—36
4
Go & 303
0—36
2
64
0—32
1
7'
0—32
3
77
59
1
80
40—50
99
16—23
105
275
1
116
'T "t
1 ~
1
"7
SO
'33
0—36
144
45
3
156
469
1
164
32
5
204
75—94
2
213
0—36
2
240
9—45
2
250
20—45
1
257
0 — 52
1
273
13
3
274
57
4
282
27—54
i
305
113
2
310
73
1
3-1
82
1
4
6
3
2
1
2
2
9
1
3
2
1
6
2
1
3
1
1
4
5
2
9
1
2
1
6
3
5
1
3
7
1
2
3
1
4
1
7
1
15
42
14
37
65
31
52
3i
6
17
17
54
O ->
"J
87
13
44
33
15
24
65
40
26
18
55
12
The last column in the above table indicates the total number of species of Polyzoa found at the
respective Stations; and it therefore includes the numbers given in the preceding three columns, in addition
to that of the Cheilostomata, which are throughout more numerous than the other three groups together.
As the revision of the Cheilostomata has not yet been completed, the numbers in the last column are
probably not quite accurate.
168
169
or with both these groups; with the total number of species, in each of the three groups, found
at each Station, foliowed by the total numbers of species of Polyzoa recorded. The table includes
those Stations which were most productive for Polyzoa in general ; the highest figures being
reached by Stat. 164, with 87 species, and Stations 77 and 273, with 65 species each. It will
be seen, by a simple process of subtraction, that the Cheilostomata are, in all these cases, in
excess — and usually largely in excess — of the other three groups together. The percentage
of Cyclostomata to the total number is, however, by no means uniform; ranging from 2.5 p.c.
at Stat. 274 (57 M.) to 29 p.c. at Stat. 105 (275 AL). This indicates that the conditions at the
various Stations are not equally favourable for the growth of Cyclostomata, although it is not
obvious why some are better than others. There is some reason to think that the Cyclostomata
are, on the average, better adapted for deep water than the Cheilostomata; and this is brought
out by the following Stations, not included in the table under consideration : — Stat. 1 19 (1901 M.),
with 50 p.c. of Cyclostomata (Cycl. 1, Cheil. 1); Stat. 97 (564 M.), with the same numbers;
Stat. 139 (397 M.), with 33.3 p.c. (Cycl. 1, Cheil. 2); and Stat. 59 (390 AL), with 25 p.c.
(Cycl. 2, Cheil. 6). But these numbers are too small to prove much; and it may be noted that
Stat. 251 (204 AL) yielded only 6.6 p.c. of Cyclostomata (Cycl. 1, Cheil. 14); while another
from shallow water (Stat. 315, o — 36 AL) gave the high proportion of 23 p.c. of Cyclostomata
(Cycl. 6, Cheil. 20).
The table on p. 168 reveals certain marked differences between the Cyclostomata and the
Ctenostomata ; but this is not surprising, in view of the fact that the Cyclostomata are strongly
calcified, whereas the Ctenostomata are non-calcareous. The largest number of Cyclostomata (9)
was yielded by Stat. 105 (275 AL), where no Ctenostomata were found. Stat. 310(73 AI), similarly
characterised by the absence of Ctenostomes, provided 7 Cyclostomes. Stat. 240 (9 — 45 AL)
had 7 Cyclostomes and 2 Ctenostomes. In certain other cases the advantage was with the
Ctenostomes, as at Stat. 164, 32 AL (Cycl. 5, Cten. 9); Stat. 273, 13 AL (3,6); Stat. 71,
o — 32 AL (1,6); and Stat. 64, o — 32 AL (1,4).
Cyclostomata occurred with Entoprocta in a larger number of Stations than with Cteno-
stomata; but it will be seen by the table on pp. 92 — 94 that a considerable proportion of the
records of Entoprocta are of species of Loxosoma which live on Rctcpora and other calcareous
Cheilostomes. The table on p. 168 shows that the various Stations are not equally favourable
for the occurrence of Cyclostomata and Entoprocta. This may be illustrated by Stat. 105, 275 AL
(Cycl. 9, Ent. 1), Stat. 282, 27— 54 AL (4,1), Stat. 274, 57 AL and Stat. 53, o— 36 AL (1,4) and
Stat. 164, 32 AL (5,5). Here again, no satisfactory explanation of the differences can be given.
The majority of the Cyclostomes obtained came from shallow water. Two thirds ot the
records (65 out of a total of 98) were from depths less than 60 Aletres. Of the remaining
33 specimens, 14 were dredged in depths from 69 to 94 AL; 1 from 113 AL; and 18 trom
204 to 1901 AL The greatest depth (Stat. 119, 1901 AL) at which Cyclostomes were obtained
yielded only one species, Tubulipora cassiformis, obtained also at Stat. 156, 469 AL This may
accordingly be regarded as an abyssal form. Hornera spinigera was obtained in three deep
dredgings (Stations 97, 156, 105; 564, 469, 275 AL) but also at Stat. 117, So Aletres. The
single fragment of H. caespitosa came from Stat. 105, 275 AL; and the genus Hornera is thus
169
SIBOGA-EXPEDITIE XXVIII a. 22
170
al most confined to deep water, so far as the 'Siboga' dredgings are concerned. Titbulipora
atlantica, var. flextiosa was obtained only at Stations 156 and 105, 469 and 275 M. Of the
two species of Berenicea, B. lineata was obtained once at Stat. 139, 397 M., but once at only
9 — 45 M. (Stat. 240); while B. sarniensis occurred at Stations 105, 204 and 310, (275, 75 — 94
and 73 M.). Entalophora proboscidea ranged from 390 M. to some depth less than 32 M., the
determination of the deep water specimen being doubtful. The specimens referred to Tubulipora
pulcherrima were numerous, and are recorded from 390 M. to the shallowest dredgings; four
of the records being, however, from more than 100 M., and three from more than 200 M.
Cr is ia elongata and three species of Lichcnopora are each recorded from Stat. 105, 275 M.,
although all the other occurrences were from less than 95 M. None of the other Cyclostomes
found were from depths exceeding 91 M.
If the determinations made are correct, it thus appears that several of the species range
from quite shallow water to depths that can be considered abyssal. But it is not impossible
that this apparently wide range may be due to incorrect determinations of the specimens from
deep water.
With regard to the Geographical Distribution of the species in the Archipelago, it may
be pointed out that the Stations where Cyclostomes were found are distributed over the whole
of the area investigated. The Sulu Sea, which yielded no Entoprocta and only one Ctenostome,
has furnished 9 Cyclostomes. Crisina radians was found only at two Stations (37, 315), close
together, off the Paternoster Islands; but Crisict elongata and Tubulipora pulcherrima, the
species of which the records are most numerous, were widely distributed in the Archipelago.
It thus appears that the area investigated by the 'Siboga' cannot be divided into distinct sub-
regions, on the evidence of the Cyclostomata, any more than was the case with the Entoprocta
or the Ctenostomata.
170
BIBLIOGRAPHY ').
AUDOUIN, V., 1826, "Explication sommaire des Planches de Polypes de 1'Égypte", in SAVIGNY's "Description
de 1'Égypte" (q. v.). For date of publication, see SHERBORN, C. D., Proc. Zool. Soc. 1897, p. 287.
1828, Ibid., 2e Édition, T. XXIII, p. 40.
Blainville, H. M. D. de, 1834, "Manuel d'Actinologie".
BUSK, G., 1S75, "Catalogue of Marine Polyzoa in the British Museum", Part III, "Cyclostomata".
1SS6, "Report on the Polyzoa (Part II)", Challenger Reports, Pt L.
CALVET, L., 1907, "Bryozoaires", Expéd. Sci. 'Travailleur' et 'Talisman', T. VIII.
Grav, J. E., 1848, "List of the Specimens of British Animals in the British Museum", Pt I, "Centroniae
or Radiated Animals".
HARMER, S. F., 1891, "On the British Species of Crisia", Quart. J. Mier. Sci., XXXII, p. 127.
1896, "On the Development of Llchenopora verrucaria Fabr.", Ibid., XXXIX, p. 71.
HlNCKS, T., 1880, "History of the British Marine Polyzoa".
■ 1884, "Contr. Gen. Hist. Mar. Pol.", XII, "Polyzoa from India (Coast of Burmah)", Ann. Mag. Xat.
Hist. (5) XIII, p. 356.
!887, "Polyzoa and Hydroida of the Mergui Archipelago", J. Linn. Soc. Zool. XXI, p. 121.
JELLY, E. C, 1S89, "A Synonymie Catalogue of the Recent Marine Bryozoa".
JOHNSTON, G., 1838, "A History of the British Zoophytes", (ist Ed.).
1847, Ibid., 2Dd Ed.
JULLIEN, J., 1888, "Mission Scientifique du Cap Horn, 1882— 1883", T. VI, Zool., "Bryozoaires".
and CALVET, L., 1903, "Bryozoaires provenant des Campagnes de 1'Hirondelle", Res. Camp. Sci.
Albert ier Prince de Monaco, Fase. XXIII.
KlRKPATRICK, R., 1888, "Polyzoa of Mauritius", Ann. Mag. Nat. Hist. (6) I, p. 72.
18901, "Report upon the Hydrozoa and Polyzoa.. China Sea", Ann. Mag. Nat. Hist. (6) V, p. 11.
1S902, "Reports on the Zoological Collections made in Torres Straits by . . A. C. Haddüx", "Hydroida
and Polyzoa", Proc. R. Dublin Soc. (N. S.) VI, p. 603.
LAMARCK, J. B. P. A. DE, 18 16, "Histoire Naturelle des Animaux sans Vertèbres", T. II.
Ibid., 2e Édition, T. II, par G. P. Deshayes et H. Milne Edwards.
1) Full references to most of the Memoirs cited are given in the text. The present list includes only those works which are
quoted in several places where the full bibliographic reference is not given.
171
LaMOUROÜX, J. V. F., 1816, "Histoire des Polypiers Coralligènes Flexibles".
1S21, "Exposition Méthodique des Genres de 1'Ordre des Polypiers".
. 1824, 1827, in "Encyclopédie Méthodique", "Histoire Naturelle des Zoophytes, ou Animaux rayonnés".
ivr. 95, 1824. pp. 1—37 cf Sherborn c D- and Woodward, B. Ii., Proc. Zool. Soc, 1899, p. 595.
I.ivr. 9S, 1827, pp. 377—819 )
LEVINSEN, G. M. R., 1894, "Mosdyr", Zool. Danica, 9 Hefte.
NlCKLES, 1. M. and Bassler, R. S., 1900, "A Synopsis of American Fossil Bryozoa", Buil. U. S. Geol.
Survey, N" 173 (valuable for Bibliography of Polyzoa, recent and fossil).
D'OrbigNV, A., 1851 — 1854, "Paleontologie Franchise", "Terrains Crétacés", T. V, "Bryozoaires". According
to C. D. Sherborn (Geol. Mag., Dec. IV, Vol. VI, 1899, p. 224, the dates of publication of this
work were as follows : —
Livr. 165 — 176, pp. 1— 188; — 1S51.
Livr. 177—194, pp. 189—472; — 1852.
Livr. 195—214, pp. 473—984; — 1853.
Livr. 215 — 21S, pp. 985 — 1192; — 1854.
Ortmanx, A., 1889, "Die Japanische Bryozoenfauna", Arch. f. Naturg., Jahrg. LVI, Bd I, p. 1.
OSBl'RN, R. C, 1912, "The Bryozoa of the Woods Hole Region", Buil. Bureau Fisheries, XXX, 1910, p. 205.
PHILIPPS, E. G., 1899, "Report on the Polyzoa", WlLLEY's Zool. Results, Pt IV.
ROBERTSON, A., 1910, "The Cyclostomatous Bryozoa of the West Coast of North America", Univ. California
Publ., Zool., VI, N° 12, p. 225.
SAVIGNV, J. C, "Description de 1'Égypte" (Plates only): — see AUDOUIN.
Tiiornelv, L. R., 1905, "Polyzoa", W. A. Herdman's Report to the Government of Ceylon on the Pearl
Oyster Fisheries of the Gulf of Manaar (published by the Royal Society), Part IV, "Suppl. Rep.",
XXVI, p. 107. See also 1906, "Corrections and Additions", Ibid., Pt V, Preface, p. 449.
. 1907, "Report on the Marine Polyzoa in the . . Indian Museum", Ree. Ind. Mus., I, p. 179.
I9I2, "The Marine Polyzoa of the Indian Ocean, from H. M. S. Sealark", Trans. Linn. Soc, 2 Ser.
Zool., XV, p. 137.
THURSTON, E., 1895, "Ramésvaram Island and Fauna of the Gulf of Manaar", Madras Government Museum,
Buil. N° 3, 2«d Ed. (pp. 131, 132; List of Polyzoa, determined by R. KlRKPATRICK).
Waters, A. W., 1904, "Bryozoa", Expéd. Antarct. Beige, Rés. Voy. S. Y. Belgica", Zoölogie.
1^09, "Reports on the Marine Biology of the Sudanese Red Sea", XII, "The Bryozoa", Pt I,
"Cheilostomata", J. Linn. Soc. Zool., XXXI, p. 123.
1910, Ibid., Pt II, "Cyclostomata, Ctenostomata, and Endoprocta", t. cit., p. 231.
1913, "The Marine Fauna of British East Africa and Zanzibar", "Bryozoa Cheilostomata", Proc.
Zool. Soc, p. 458
1914, Ibid., "Bryozoa Cyclostomata, Ctenostomata, and Endoprocta", Proc. Zool. Soc. p. 831.
WHITELEGGE, T., 1889 (1890), "List of the Marine and Fresh-water Invertebrate Fauna of Port Jackson
and Neighbourhood", J. Proc. R. Soc. N. S. Wales, XXIII, pp. 282—293, 322, 333.
WOODWARD, B. B., 1903— 1915, "Catalogue of the Books, Manuscripts, Maps and Drawings in the British
Museum (Natural History)", Vols I— V. (This catalogue contains valuable information with regard
to the Dates of Publication of many of the Memoirs cited).
172
INDEX.
acervata, Amathia, 68, 70.
acropora, Crisia, 97, 98 n.
alatum, Loxosoma, 7, 8.
albidum, Alcyonidium, 56.
Alcyonella, 35.
„ articulata, 35.
Alcyonellea, 35.
Alcyonidiées, 36.
Alcyonidiidae, 36.
Alcyonidium, 35, 36, 38;
distribution, 36,95.
„ albidum, 56.
„ antarcticum, 36.
„ cellarioides, 56.
„ diaphanum, 36.
„ flabelliforme, 36.
„ flustroides, 36.
„ gelatinosum, 36, 38.
mytili, 36, 37.
„ pedunculatum, 36.
polyoum, 37, 95.
Alcyonidulae, 36.
altum, Cylindroecium, 57 11.
Amathia, 43, 60, 64; distribution,
92—95.
„ acervata, 68, 70.
„ brasiliensis, 70.
„ convoluta, 64, 67, 69, 95,
„ crispa, 64, 68.
„ distans, 68, 69.
„ pruvoti, 69.
„ semiconvoluta, 65.
semispiralis, 64, 66, 6y.
„ spiralis, 64, 67.
„ tortuosa, 64, 6S, 69.
„ Loxosoma on, 5.
annectens, Nolella, 52, 55, 57.
annelidicola, Loxosoma, 14, 23.
annulata, Berenicea, 114.
annulatum, Loxosoma, 5, 11.
antarcticum, Alcyonidium, 36.
Antedon, Loxosoma on, 9.
antedonis, Loxosoma, 9, 15.
aperta, Tubulipora, 122, 124, 132,
143 n.
aperture, a part of the frontal
sitrface closed by a delicate mem-
brane, 85, 86, 88.
Apsendesia, 155.
„ cristata, 155.
Arachnidia, 48, 49.
„ hippothoöides, 48.
Arachnidiidae, 43, 48, 52.
Arachnidium, 43, 48, 50, 52;
distribution,^},, 96.
„ fibrosum, 50, 51.
„ irregulare, 49, 50,96.
„ simplex, 86.
Arachnoidea, 43, 48, 50, 52;
distribution, 92, 96.
„ protecta, 50, 96.
„ ray-lankesteri,50, 51.
Arachnoidia, 50.
armata, Buskia, 88.
„ , Hippuraria, 88, 92.
„ , Vesicularia, 88.
Arthropodaria, 25.
„ benedeni, 26.
articulata, Alcyonella, 35.
„ , Paludicella, 35, 56.
Ascopodaria, 25.
„ discreta, 29.
„ gracilis, 28.
„ misakiensis, 29.
„ nodosa, 28.
Ascorhiza occidentalis, 36.
asteriscus, Hypocytis, 153.
atlantica, Farella, 73, 84.
atlantica, Farrella,- 73, 74, 84, 85.
„ Idmonea, 122, 124, 139.
v Tubulipora, 124, 138.
v Valkeria, 46, 73, 84, 94,
96.
r var. disticha, Idmonea,
124.
„ forma erecta, Tubuli-
pora, 125.
v var. flexuosa, Tubuli-
pora, 127, 170.
„ var. tenuis, Idmonea,
127, 12S.
australis, Barentsia, 24 n.
„ , Buskia, S6.
„ , Idmonea, 120, 121, 12511.
„ , Pedicellina, 24, 25,29, 34.
Avenella, 43, 60.
Barentsia,
23> 25; distribution,
92—95.
ï!
australis, 24 n.
V
belgica, 26.
)!
benedeni, 26.
bulbosa, 25, 26.
capitata, 32 n.
T
discreta, 29, 34.
1}
geniculata, 33.
V
gracilis, 26, 27, 28, 34.
V
laxa, 32.
V
macropus, 30, 32.
V
major, 32 11.
V
„ , var. elongata,
32 n.
TT
misakiensis, 29, 30.
TT
nodosa, 28.
TT
ramosa, 30.
TT
timida, 29, 32.
r
variabilis, 32 n.
'74
Harentsia variarticulata. 25, 26.
sp., 32 n.
Beania, 60.
belgica, Barentsia, 26.
benedeni, Arthropodaria, 26.
, Barentsia, 26.
bengalensis, Victorella, 44, 47, 48.
Berenicea, 114, 118, 123; distri-
bution, 166, 167, 170.
„ annulata, 1 14.
„ fasciculata, 123.
„ lineata, 115, 116, 123,
170.
„ patina, 117.
„ prominens, 114.
„ sarniensis, 114, 117 n.,
170.
„ suborbicularis, 116.
biciliata, Crisia, 103.
bicolor, Diastopora, 116.
„ , Liripora, 1 16, 1 17.
bigeminata, Mimosella, 79, 82.
bilateralis, Vesicularia, 70.
binderi, Flustrella, 40.
biserialis, Bowerbankia, 70.
blainvillii, Elzerina, 38, 39, 95-
„ , Farciminaria, 39.
boeckii, Triticella, go, 91.
boryi, Proboscina, 1 19.
Bowerbankia, 35, 43, 60, 62 n., 70 ;
distribution, 92 — 93.
„ biserialis, 70.
„ caudata, 72.
B densa, 71.
„ imbricata, 6r, 70, 72.
„ „ , ƒ<?/'/« den-
sa, 71.
„ pustulosa, 61, 72.
brasiliensis, Amathia, 70.
breusingi, Pedicellina, 25.
breve, Loxosoma, 5, 19.
brongniartii, Chorizopora, 1 14.
brumpti, Loxosoma, 9, 10, 15.
Bryozoa, 4.
bulbosa, Barentsia, 25, 26.
Buskea, 85.
buski, Lichenopora, 161.
Buskia, 43, 85; distribution, 92 — 96.
„ armata, 88.
„ australis, 86.
„ nitens, 85.
„ pilosa, 89.
setigera, 43, 85, 87, 89, 94,
96.
Buskia socialis, 43, 85, 86.
Buskiidae, 85.
caespitosa, Hornera, 149, 150, 169.
californica, Lichenopora, 161.
„ , Unicavea, 16 [.
i Calyx, the part of the body contain-
ing the viscera, in Entoprocta.
Caiupylonemida, 46, 73.
canaux de r en f or cement, 126, 13S,
145 n-
Cancelli , cavities resembling the
initial stages of zooecia, special ly
characteristic of Lichenoporidae.
Canda, Loxosoma on, 5, 21.
capitata, Barentsia, 32 n.
Camosa, 35.
cassiformis, Tubulipora, 135, 169.
caudata, Bowerbankia, 72.
Caveidae, 153.
cellarioides, Alcyonidium, 56.
Cellepora, Loxosoma on, 5, 19.
cenomana, Crisina, 138.
„ , Crisisina, 138.
„ , Idmonea, 138.
Cephalodiscus, Loxocalyx on, 5, 8.
cernua, Pedicellina, 31.
Cheilostomata, distribution, 168, 169.
Chorizopora brongniartii, 114.
ciliata, Discopora, 161, 164.
„ , Discoporella, 161, 164.
, Lichenopora, 161, 162, 164.
circinata, Crisia, 103, 105.
circulare, Loxosoma, 5, 16, 17.
circumplicata, Sirinx, 64, 68.
cirriferum, Loxosoma, 5, 12, 14,
15, 16.
Clavopora hystricis, 36.
closure, of Cyclostomes, 115.
clypeata, Defrancia, 155.
, Pelagia, 155.
cocciforme, Loxosoma, 5, 22, 94.
cochlear, Loxosoma, 7, 8.
Collections reported on, 3.
colonialis, Loxosomatoides, 22 n.
compacta, Pedicellina, 24.
concentrica, Flustrella, 40.
concinna, Tubulipora, 123.
Conescharellina, Loxosoma on, 5,
19.
connata, Tubulipora, 123.
continentalis, Victorella, 44,47,48.
convoluta, Amathia, 64, 67, 69, 95.
„ , Serialaria, 6j.
i74
Coppingeria, Loxosoma on, 4.
Corallina, 154.
Cordyle, 90.
Cornea, 35.
cornuta, Crisia, 107.
„ , var. geniculata, Crisia, 106.
» » » 1 Crisidia,
106.
crassicauda, Loxosoma, 4, 15.
„ , Loxosomella, 6.
cribraria, Crisia, 10 1 n.
Crisia, 96 ; distribution, 1 66, 1 67, 1 70.
„ acropora, 97, 98 n.
„ biciliata, 103.
„ circinata, 103, 105.
„ cornuta, 107.
„ „ ,var. geniculata, 106.
„ cribraria, 101 n.
„ cuneata, 103, 106.
„ cylindrica, 103, 104.
„ denticulata, 97, 98, 99, 100,
ior, 102.
„ „ , var. gracilis, 97,
101.
„ eburnea, 101 n.
„ „ , var. laxa, 105.
„ edwardsiana, 103.
„ elongata, g6, 104, 170.
„ geniculata, 105. 106.
„ hamifera, 9S n.
„ holdsworthii, 97, 98.
„ hörnesii, 98.
„ inflata, 103.
„ kerguelensis, 105-
„ margaritacea, 97.
„ nigrijuncta, 97, 102.
„ occidentalis, 101 n.
„ patagonica, 101.
„ punctifera, 97.
„ sinensis, 97.
„ subaequalis, 98.
„ terrae-reginae, 97, 10 1.
Crisiadae, 96.
Crisidia cornuta, var. geniculata,
106.
Crisiidae, g6.
Crisina, 122, 137, 138, 139;
distribution, 166, 167, 170.
„ cenomana, 138.
„ hochstetteriana, 138.
, Ligeriensis, 138.
, Normaniana, 137.
„ radians, 123, 126, 137, 138,
139- l7°-
175
Crisina ramosa, 137.
„ subgracilis, 13S.
subgradata, 137.
„ triangularis, 137, 138.
„ unipora, 137.
Crisisina, 137, 138.
„ cenomana, 13S.
crispa, Amathia, 64, 68.
„ , Serialaria, 64.
cristata, Apsendesia, 155.
„ , Radiopora, 160 n.
Cruciform Stolonifera, 43.
Cryptocyst, a calcareous, iris-hke
shelf, developed beneath tin- mem-
brane covering the f rental sitr-
face of a zooecium or canccllus.
Cryptopolyzoon, 43.
Ctenostomata, 35; distribution,<$2 —
96, 168 — 170.
cuneata, Crisia, 103, 106.
Cupularia umbellata, 155.
cuscuta, Yalkeria, 61.
„ , Vesicularia, 61.
Cuscutaria verticillata, 83.
Cyclostomata, 96; distribution, 166-
170.
cylindrica, Crisia, 103, 104.
„ ,var. <?ƒ Flustrella hispida,
40.
Cylindroeciidae, 43, 52.
Cylindroecium, 43, 52, 53, 54.
„ altum, 57 11.
„ dilatatum, 53, 54.
„ giganteum, 53, 54.
papuense, 53.
Cytisidae, 150.
davenporti, Loxosoma, 14.
Dedalaea mauritiana, 70.
deflexa, Entalophora, III, 112.
Defïancia, 155.
„ clypeata, 155.
„ lucernaria, 153.
delicatula, Entalophora, 110, III,
1 13-
fl , Pustulipora, I 10.
„ , Pustulopora, 1 10.
densa, Bowerbankia, 71.
„ , var. of Bowerbankia imbri-
cata, 71.
denticulata, Crisia, 97, 98, 99, 100,
101, 102
„ , var. gracilis, Crisia, 97,
101.
diaphanum, Alcyonidium, 36.
Diastopora, 114, 11S, 119.
„ bicolor, 116.
„ fasciculata, 123.
„ lincata, 116, 123.
B prominens, 1 16.
„ sarniensis, 114, II 5.
„ „ , var. peran-
gusta, 115.
Diastoporidac, 113.
dichotoma, Farciminaria, 39.
„ , Flustrella, 39, 40.
„ , Verrucularia, 39, 40.
digitata, Fasciculipora, 152.
„ , Supercytis, 151, 152, 153.
dilatata, Nolella, 59.
dilatatum, Cylindroecium, 53, 54.
Discocavea, 154, 155.
„ irregularis, 155 n.
Discopora, 11 8, 119.
„ ciliata, 161, 164.
„ hispida, 1 19, 154.
Discoporella, 153, 154, 155.
„ ciliata, 161, 164.
„ holdsworthii, 155,
160, 161.
„ mediterranea, 164, 165.
„ novae-zelandiae, 155.
„ porosa, 1 64.
„ umbellata, 155-
Discotubigera lineata, 116.
discreta, Ascopodaria, 29.
„ , Barentsia, 29.
, Tervia, 143.
Disporella, 153, 154.
„ hispida, 154.
distans, Amathia, 68, 69.
disticha, var. 0/Tdmonea atlantica,
124.
eburnea, Crisia, 101 n.
„ , var. laxa, Crisia, 105.
echinata, Lichenopora, 160, 161.
„ , Pedicellina, 33 n.
Echinella placoides, 40 n.
Ectoprocta, 35.
cdwardsiana, Crisia, 103.
egertoni, Hippuraria, 83, 90, 91,92.
„ , Triticella, 90, 91.
elongata, Crisia, g6, 104, 170.
„ , Hippuraria, 92.
„ , Triticella, 92.
v , var. of Barentsia major,
32 n.
175
Elzerina, 38, 40, 41 ; distribittion, 95.
„ blainvillii, 38, 39, 95.
Entalophora, 107, 119, 144; distri-
bittion, 166, 167, 170.
„ deflexa, 1 1 1, 1 12, 1 13.
delicatula, HO, III.
fragilis, 1 1 1 .
r indica, 10S.
, intricaria, 112.
„ proboscidea,lo8, 1 1 1 ,
170.
v proboscidioides, 110.
r raripora, 108 n.
„ virgula, 108.
, wasinensis, 1 1 1, 112.
Entaloplioridac, 107.
Entaloplioridca, 107.
Entoprocta, 4, 27; distribution, 92-
96, 16S — 170.
erecta, var. of Tubulipora atlantica,
125.
excretory vesicles, 78, 132.
expansa, Hypophorella, 56.
familiaris, Plumatella, 90.
Farciminaria, 40.
„ blainvillii, 39.
„ dichotoma, 39.
Farella atlantica, j^-
Farrclla, 35, 43, 60, 75.
„ atlantica, 73, 74, 84, 85.
„ fusca, 55 n.
gigantea, 53.
„ repens, 75, 80 n.
fasciculata, Berenicea, 123.
„ , Diastopora, 123.
„ , Liripora, 123.
Fasciculipora, 150.
„ digitata, 152.
fibrosum, Arachnidium, 50, 51.
Filicrisia geniculata, 106.
Filisparsa, 143.
, tubulosa, 143.
r varians, 144.
„ violacea, var. tubulosa,
fissurata, Hornera, 138.
flabellaris, Flustrella, 40 {misprinted
flabellaria).
„ , Phalangella, 123.
v , Pherusa, 40.
v , Tubulipora, 121, 122.
flabelliforme, Alcyonidium, 36.
flava, Triticella, 90.
r76
flexuosa, Idmonea, 127, 12S.
, var. of Tubulipora atlan-
tica, 127, 170.
Flustra, Loxosoma on, 20 n.
Flustrella, 40, 41.
binderi, 40.
„ concentrica, 40.
, dichotoma, 39, 40.
flabellaris, 40 (misprinted
flabellaria).
hispida, 39, 40, 41, 56.
„ „ , var. cylindrica,
40.
Flustrellidae, 38.
flustroides, Alcyonidium, 36.
folini, Tervia, 143.
Folliculina, occurring with Loxo-
soma, iS.
fragilis, Entalophora, III.
„ , Pustulipora, III.
frondiculata, Hornera, 150.
Frondipora verrucosa, 165.
fruticosa, Pedicellinopsis, 26, 30.
fusca, Farrella, 55 n.
„ , Vesicularia, 55 n.
Galeidae, 153.
gelatinosum, Alcyonidium, 36, 38.
geniculata, Barentsia, 33.
„ , Crisia, 105, 106.
„ , Filicrisia, 106.
„ , var. of Crisia (Crisidia)
cornuta, 106.
gigantea, Farrella, 53.
, Nolella, 54.
giganteum, Cylindroecium, 53, 54.
Gonypodaria, 25, 26.
, nodosa, 28.
n ramosa, 25, 26, 30.
gracilis, Ascopodaria, 28.
, Barentsia, 26, 27, 28, 34.
„ , Alimosella, 78 n., 79, 80 n.,
Si.
, Pedicellina, 27.
„ , Urnatella, 26.
„ , var. nodosa, Pedicellina, 28.
„ , var. of Crisia denticulata,
97. IOI.
gracillima, Idmonea, 12S.
granulata, Stomatopora, 120.
Gytnnolaemata, 35.
Halichondria, Loxocalyx on, 5, 6.
Halcyonella, 35.
Halcyonellea, 35.
hamifera, Crisia, 98 n.
harmeri, Loxosoma, 15.
hippothöoides, Arachnidia, 4S.
Hippuraria, 83, 88, 90, 91, 92.
„ armata, 88, 92.
egertoni, 83, 90, 91,92.
„ elongata, 92.
„ verticillata, 81, 85,92.
Hislopia, 40 n.
Hislopiidae, 48.
hispida, Discopora, 119, 154.
„ , Disporella, 154.
, Flustrella, 39, 40, 41, 56.
„ , Lichenopora, 119, 154,
162, 164.
„ , var. cylindrica, Flustrella,
40.
hochstetteriana, Crisina, 138.
holdsworthii, Crisia, 97, 98.
, , Discoporella, 155,
160, 161.
„ , Lichenopora, 155,
161.
Homodiaetidae, 44.
Hornera, 147; distribution, 166,
167, 169.
„ caespitosa, 149, 150, 169.
„ fissurata, 138.
„ frondiculata, 150.
spinigera, 147, 169.
„ tubulosa, 143.
„ violacea, var. tubulosa,
143-
Horneridae, 147.
hörnesii, Crisia, 98.
Hypocytis asteriscus, 153.
Hypophorella, 43, 80 n.
, expansa, 56.
hystricis, Clavopora, 36 n.
Idmonea, 119, 122, 123, 126, 137,
141, 143.
„ atlantica, 122, 124, 139.
„ „ , var. disticha,
124.
„ „ , var. tennis,
127, 128.
„ australis, 120, 121, 125 n.
„ cenomana, 138.
„ flexuosa, 127, 128.
„ gracillima, 128.
„ interjuncta, 129, 130,
132 n., 134.
176
Idmonea irregularis, 143, 144, 146.
„ milneana, 129, 130, 132 n.,
133. '34-
„ pedata, 131 n.
„ pulcherrima, 129, 132.
radians, 123,125,138, 139.
„ radicata, 129, 131 n., 134.
„ ramosa, 120, 137, 138.
„ serpens, 122.
„ subgracilis, 138.
„ tenella, 127, 129.
„ tortuosa, 126 n.
n triquetra, 122.
„ tuberosa, 129.
unipora, 137, 138.
Idmoneidae, 107, 11 8.
Idmoniidae, 123.
imbricata, Bowerbankia, 61, 70, 72.
„ , Sertularia, 70.
„ , Vesicularia, 61.
„ , form densa, Bowerban-
kia, 71.
imperialis, Lichenopora, 165 n.
indica, Entalophora, 108.
inflata, Crisia, 103.
interjuncta, Idmonea, 129, 130,
132 n., 134.
„ , Tubulipora, 132, 133,
'34-
intricaria, Entalophora, 112.
„ , Pustulopora, 112.
investigatoris, Platypolyzoon, 51.
irregulare, Arachnidium, 49,50,96.
irregularis, Uiscocavea, 155 n.
„ , Idmonea, 143, 144, 146.
, Tervia, 143, 144, 146.
jellyae, Tervia, 143.
Kamptoderm, the invaginablc part
of the body-wall.
kefersteinii, Loxosoma, 5, 12.
kenozooecia, 1 30.
kerguelensis, Crisia, 105-
korenii, Triticella, 90, 92.
Lagenella, 75 n.
„ nutans, So n., 81.
lanchesteri, Loxosoma, 5, 8.
laxa, Barentsia, 32.
„ , var. of Crisia eburnea, 105.
Lepralia, Loxosoma on, 5, 17, 18.
leptoclini, Loxocalyx, 5, 8.
„ , Loxosoma, 7, 8.
/ /
Lichenopora, 153, 154, 160; distri-
bution, 166, 167,
170.
„ buski, 161.
„ californica, 161.
„ ciliata, 161, 162, 164.
n echinata, 160.
B hispida, 119, 154,
162, 164.
„ holdsworthii, 155,
161.
„ imperialis, 165 n.
„ mediterranea, 164.
„ novae-zealandiae, 155,
156.
„ novae-zelandiae, 154,
155, 156, 160, 161,
163, 164, 165.
„ radiata, 154, 160.
reticulata, 160.
„ turbinata, 154.
„ verrucaria, 154, 156,
157, 162, 164.
„ victoriensis, 155, 160.
wanganuiensis, 164.
Lichenoporidae, 153 .
Ligeriensis, Crisina, 138.
liliacea, Tubulipora, 122, 123.
lineata, Berenicea, 115, 116, 123,
170.
„ , Diastopora, 116, 123.
„ , Discotubigera, 116.
a , Liripora, 1 16, 123.
lineatus, Loxocalyx, 5, 6.
Liripora, 117, 123.
v bicolor, 116, 117.
„ fasciculata, 123.
„ lineata, 1 16, 123.
loricatum, Loxosoma, 5, 21.
loxalinum, Loxosoma, 15.
Loxocalyx, 5, 6; distribution, 92,
93-
„ alatus, 7, S.
„ cochlear, 7, 8.
leptoclini, 5, 7, 8.
„ lineatus, 5, 6.
„ neapolitanus, 7, 8.
pes, 7, S.
„ raja, 6, 7.
„ tethyae, 7.
Loxosoma, a/iosts" of, 2, 4; distri-
bution, 92 — 95.
„ , list of species found, 5 .
„ , genus, s. str., 6, 8.
SIEOGA-EXPEDITIE XXVIII a.
Loxosoma alatum, 7, 8.
„ annelidicola, 14, 23.
„ annulatum, 5, 11.
„ antedonis, 9, 15.
„ breve, 5, 19.
„ brumpti, 9, 10, 15.
„ circulare, 5, 16, 17.
, cirriferum, 5, 12, 14,
15, 16.
„ cocciforme, 5, 22, 94.
., cochlear, 7, 8.
, crassicauda, 4, 15.
„ davenporti, 14.
„ harmeri, 15.
„ kefersteinii, 5, 12.
„ lanchesteri, 5, 8.
„ leptoclini, 7, 8.
v loricatum, 5, 21.
„ loxalinum, 15.
„ minutum, 9, 10.
„ murmanicum, 9, 10.
„ neapolitanum, 7, 8.
„ nitschei, 20.
pes, 7, 8.
„ phascolosomatum,9, 10,
11, 15.
„ pusillum, 5, 16.
raja, 7.
„ saltans, 15.
B singulare, 6, 19, 19 n.,
23
„ sluiteri, 5, 9, 17, 94.
„ subsessile, 5, ig.
„ tethyae, 7.
„ troglodytes, 5, 17, 19.
„ velatum, 5, 13.
Loxosoma, sp., on Amathia, 5.
„ , , „ Coppingeria, 4.
„ 1 Ti » Phascolion, 10,
1 1.
, , M n Phascolosoma,
4, 11.
, , „ „ Retepora, 17.
» , r, v (rgen.), ö«Xan-
thias, 5.
„ , „ from Queensland, 5.
Loxosomatidae, 4.
Loxosomatoides colonialis, 22 n.
Loxosomella, 6.
„ crassicauda, 6.
Loxosomidae, 4.
lucernaria, Defrancia, 153.
lucida, Tubulipora, 123.
Lunulites umbellata, 155.
177
macropus, Barentsia, 30, 32.
major, Barentsia, 32 n.
margaritacea, Crisia, 97.
marginata, Reptotubigera, 1 19.
mauritiana, Dedalaea, 70.
mediterranea, Discoporella, 164,
165
, Lichenopora, 164.
Melobesia, 154.
„ pustulosa, 154.
n radiata, 154, 160.
Membranipora pilosa, 56.
Menipea, Loxosoma on, 20.
Methods, 9, 41.
milneana, Idmonea, 129, 130,
132 n., 133, 134.
, , Tubulipora, 132, 133.
Mimosella, 43, 74, 78, 81 ; distri-
bution, 92, 93.
„ bigeminata, 79, 82.
„ gracilis, 79, 80 n., 81.
,, tenuis, 84.
„ verticillata, 8l, 84.
sp., S3, 84.
Mimosellidae, 78.
minutum, Loxosoma, 9, 10.
misakiensis, Ascopodaria, 29.
B , Barentsia, 29, 30.
mülleri, Paludicella, 44.
„ , Victorella, 44.
murmanicum, Loxosoma, 9, 10.
mytili, Alcyonidium, 36, 37.
neapolitanum, Loxosoma, 7, 8.
neocomiensis, Reptotubigera, 119.
nigrijuncta, Crisia, 97, 102.
nitens, Buskia, 85.
nitschei, Loxosoma, 20.
nodosa, Ascopodaria, 28.
B , Barentsia, 28.
„ , Gonypodaria, 28.
„ , var. ö/Pedice!lina gracilis,
28.
Nolella, 43 n., 52, 53, 54; distri-
bution, 92 — 96.
annectens, 52, 55, 57.
„ dilatata, 59.
gigantea, 54.
„ papuensis, 53, 94, 96.
stipata, 52, 53.
Nolellidae, 43, 52.
Xormaniana, Crisina, 137.
novae-zealandiae, Lichenopora ,
ISS, 156.
L33»
23
i;8
novae-zelandiae, Discoporella, 155.
, Lichenopora,i54>
155- '56' l6o>
161, 163, 164,
.65.
nutans, Lagenella, So 11., Si.
„ , Valkeria, Si.
, Walkeria, Si.
Obelia, iiy.
tubulifera, 119.
occidentalis, Ascorhiza, 36.
„ , Crisia, 10 1 n.
ooeciopore, 98 n.
ooeciostome, 98 n.
„ , accessory, 130, 132.
organisans, Tubulipora, 121.
c Irifice, the external opening of the
invaginated kamptoderm (q. v.).
Paludicclla, 35, 43, 44, 52> 59 "■
„ articulata, 35, 56.
„ mülleri, 44.
Paludicellea, 41, 43, 60.
Paludlcellidae, 43, 44, 52.
Paludicellina, 43.
papuense, Cylindroecium, 53.
papuensis, Nolella, 53, 94, 96.
„ , Vesicularia, 6l.
patagonica, Crisia, 10 1.
patina, Berenicea, 117.
pavida, Victorella, 44, 45.
pedata, Idmonea, 131 n.
Pedicellina, 23; distribution, 93.
australis, 24, 25, 29,
34-
„ breusingi, 25.
cernua, 31.
„ compacta, 24.
„ echinata, 33.
„ gracilis, 27.
„ „ , var. nodosa,
28.
Pedicellinea, 23 n.
Pedicellinidae, 23.
Pedicellinopsis, 25, 26.
fruticosa, 26, 30.
pedunculatum, Alcyonidium, 36.
Pelagia, 155.
clypeata, 155.
pellucidum, Zoobotryon, 70.
perangusta, var. ^/Diastopora sar-
niensis, 115.
Peristome, 48 n.
pes, Loxosoma, 7, 8.
phalangea, Phalangella, 123.
, Tubulipora, 122, 142.
Phalangella, 122.
„ flabellaris, 123.
,, phalangea, 123.
Phascolion, Loxosoma on, 5, 9, 10,
1 1.
Phascolosoma, Loxosoma on, 4, 5,
8, 9, 10, 1 1.
phascolosomatum, Loxosoma, 9, 10,
n, 15.
Pherusa flabellaris, 40.
„ tubulosa, 56.
philippsae, Reptotubigera, 120.
pilosa, Buskia, 8g.
„ , Membranipora, 56.
placoides, Echinella, 40 n.
Platypolyzoon investigatoris, 51.
Plumatella familiaris, 90.
polyoum, Alcyonidium, 37, 95.
„ , Sarcochitum, 37.
Polypide, 26.
Polyzoa, 4.
porosa, Discoporella, 164.
Pottsiella, 43.
proboscidea, Entalophora, 108, 1 1 1 ,
170.
„ Pustulipora, 108.
„ Pustulopora, 108.
proboscidioides, Entalophora, 110.
Proboscina, 1 19.
,, boryi, 119.
prominens, Berenicea, 1 14.
„ , Diastopora, 116.
protecta, Arachnoidea, 50, 96.
pruvoti, Amathia, 69.
pulcherrima, Idmonea, 129, 132.
„ , Tubulipora, 12Q, 170.
pulchra, Tubulipora, 123.
punctifera, Crisia, 97.
pusillum, Loxosoma, 5, 16.
Pustulipora, 10S.
„ delicatula, 110.
„ fragilis, 111.
„ proboscidea, 10S.
Pustuliporidae, 107.
Pustulopora, 108.
„ delicatula, 1 10.
„ intricaria, 1 12.
„ proboscidea, 10S.
pustulosa, Bowerbankia, 61, 72.
„ , Melobesia, 154.
„ , Vesicularia, 61.
17S
radians, Crisina, 123, 126, 137,
13S, 139. 170.
„ , Idmonea, 123, 125, 13S,
139-
radians, Retepora, 139.
radiata, Lichenopora, 154, 160.
„ , Melobesia, 154, 160.
radicata, Idmonea, 129, 131 n.,
L34-
Radiopora cristata, 160 n.
raja, Loxocalyx, 6.
„ , Loxosoma, 7.
ramosa, Barentsia, 30.
„ , Crisina, 137.
„ , Gonypodaria, 25, 26, 30.
„ , Idmonea, 120, 137, 138.
„ , Reptotubigera, 119.
raripora, Entalophora, 108 n.
ray-lankesteri, Arachnoidea, 50,51.
repens, Farrella, 75, 80 n.
Reptotubigera, lig, 123, 144;
distribution, 167.
„ marginata, 1 19.
„ neocomiensis, 119.
„ philippsae, 120.
„ ramosa, 1 19.
„ serpens, 119.
Retepora, Loxosoma on, 5, 11, 12,
13, 14, 16, 17.
„ radians, 139.
„ reticulata, 165.
reticulata, Lichenopora, 160.
„ , Retepora, 165.
saltans, Loxosoma, 15.
sarniensis, Berenicea, 114, 117 n.,
170.
„ , Diastopora, 114, 115.
„ , var. perangusta, Dias-
pora, 115.
Sarcochitum polyoum, 37.
Schizoporella, Loxosoma on, 5, 13,
19.
semiconvoluta, Amathia, 65.
semispiralis, Amathia, 64, 66, 67.
„ , Serialaria (Vesicula-
ria?), 67.
„ , Vesicularia, 67.
Serialaria, 60, 64.
„ convoluta, 6j.
r crispa, 64.
„ semispiralis, 67.
serpens, Idmonea, 122.
„ , Reptotubigera, 119.
179
Sertularia imbricata, 70.
„ spinosa, 61.
setigera, Buskia, 43, 85, 87,89,94,
96.
sibogae, Victorella, 45, 96.
simplex, Arachnidium, 86.
sinensis, Crisia, 97.
singulare, Loxosoma, 6, 19, 19 n.,
23-
Siphonicytara, Loxosoma on, 5, 22.
Sirinx circumplicata, 64, 68.
sluiteri, Loxosoma, 5, g, 17, 94.
socialis, Buskia, 43, 85, 86.
solida, Tervia, 143.
spinigera, Hornera, 147, 169.
spinosa, Sertularia, 61.
„ , Vesicularia, 61, 6^.
spiralis, Amathia, 64, 6j.
Spiropora, 108.
Sponges, Loxocalyx on, 7.
Stations at which were fonnd:
Polyzoa, 2, 168;
Entoprocta, 92 — 96, 168, 169;
Ctenostomata, 92 — 96, 168, 169;
Cyclostomata, 1 66 — 1 70.
stipata, Nolella, 52, 53.
Stolon, 43.
Stolonifera, 41, 43, 72.
Stomatopora, 115, 119, 120, 144.
„ granulata, 120.
subaequalis, Crisia, 98.
subgracilis, Crisina, 138.
„ , Idmonea, 138.
subgradata, Crisina, 137.
suborbicularis, Berenicea, 116.
subsessile, Loxosoma, 5> 19-
Supercytis, 150; dislribution, 167.
digitata, 151, 152, 153.
tubigera, 151.
„ watersi, 151.
symbiotica, Victorella, 44.
Syncoryne, occumng with Loxo-
soma, 12.
Synzooecipliytes, 27 n.
tenella, Idmonea, 127, 129.
tenuis, Mimosella, 84.
„ , var. of Idmonea atlantica,
127, 128,
terrae-reginae, Crisia, 97, 101.
Tervia, 123, 143, 144; dislribution,
166.
„ discreta, 143.
„ folini, 143.
Tervia irregularis, 143, 144, 146.
„ jellyae, 143.
„ solida, 143.
Tethya, Loxocalyx on, 7.
tethyae, Loxosoma (Loxocalyx), 7.
timida, Barentsia, 29, 32.
tortuosa, Amathia, 64, 68, 69.
„ , Idmonea, 126 n.
transversa, Tubulipora, 122, 123.
triangularis, Crisina, 137, 138.
trichotoma, Vesicularia, 61, 63.
triquetra, Idmonea, 122.
Triticella, 43, 88, 90.
„ boeckii, 90, 91.
„ egertoni, 90, 91.
„ elongata, 92.
flava, 90.
„ korenii, 90, 92.
Triticellidae, 83, go.
Tritilcllidae, 90.
troglodytes, Loxosoma, 5, 17, 19.
Tropics, Polyzoa in, 1.
tuberosa, Idmonea, 129.
„ , Valkeria, 76, 96.
tubigera, Supercytis, 151.
tubulilera, Obelia, 119.
Tubulipora, 118, 119,122, 123, 137,
144, 154; distribution,
166, 167, 169, 170.
,, aperta, 122, 124, 132,
143 n.
„ atlantica, 124, 13S.
„ „ , forma erecta,
125.
„ „ , var. flexuosa,
127, 170.
„ cassiformis, 135, 169.
„ concinna, 123.
„ connata, 123.
„ flabellaris, 121, 122.
„ interjuncta, 132, 133,
134-
„ liliacea, 122, 123.
„ lucida, 123.
milneana, 132, 133.
„ organisans, 121.
„ phalangea, 122, 142.
„ pulcherrima, 12g, 170.
„ pulchra, 123.
„ transversa, 122, 123.
Tubuliporidae, 118.
tubulosa, Filisparsa, 143.
„ , Hornera, 143.
„ , Pherusa, 56.
179
tubulosa, var. of Filisparsa (Hor-
nera) violacea, 143.
turbinata, Lichenopora, 1 54.
umbellata, Cupularia, 155.
„ , Discoporella, 155.
„ , Lunulites, 155.
Unicavea californica, 161.
unipora, Crisina, 137.
, Idmonea, 137, 138.
Urnatella, 26.
„ gracilis, 26.
uva, Valkeria, 75, j6, 80 n.
Valkeria, 35, 43, 60, 62 n., 73,
80 n.; distribution ,
92 — 96.
atlantica, 46, 73, 84, 94,
96.
„ cuscuta, 61.
„ nutans, Si.
„ tuberosa, 76, 96.
uva, 75, 76, 80 n.
„ verticillata, 8 1 .
Valkeriidae, 73.
variabilis, Barentsia, 32 11.
varians, Filisparsa, 144.
variarticulata, Barentsia, 25, 26.
velatum, Loxosoma, 5, 13.
verrucaria, Lichenopora, 154, 156,
157, 162, 164.
verrucosa, Frondipora, 165.
Verrucularia, 40, 41.
„ dichotoma, 39, 40.
verticillata, Cuscutaria, 83.
„ Hippuraria, 81,85,92.
Mimosella, 8l, 84.
„ Valkeria, Si.
Vesicularia, 35, 43, 60, 6l; distri-
bution, 93.
„ armata, 88.
„ bilateralis, 70.
„ cuscuta, 61.
fusca, 55 n.
„ imbricata, 61.
„ papuensis, 61.
„ pustulosa, 61.
„ semispiralis, 67.
„ spinosa, 61, 63.
„ trichotoma, 61, 63.
Vesiculariadac, 60.
( 'csiailarudac, 60.
Vesicularina, 41, 43, ÓO-
Vestibule, the distal part of the
invaginated kamptoderm (q.v.).
iSo
Victorella, 43, 44, 46, 47, 48, 52;
distribution, 92, 96.
„ bengalensis, 44, 47, 48.
continentalis, 44, 47,
48.
muilen, 44.
„ pavida, 44, 45.
sibogae, 45, 96.
„ symbiotica, 44.
Victorellidae, 43, 44.
Victorellides, 44.
victoriensis, Lichenopora, 155, 160. , wasinensis, Entalophora, 111, 112.
Vinella, 78 n. I vvatersi, Supercytis, 151.
violacea, var. tubulosa, Filisparsa,
143- Xanthias, Loxosoma (?) on, 5.
„ , var. tubulosa, Hornera, ,
143-
virgula, Entalophora, 10S.
Walkeria, 73.
„ nutans, 8r.
wanganuiensis, Lichenopora, 164. | Zooecium, 27.
Zoobotrion, 70 n.
Zoobotryon, 43, 60, 70.
„ pellucidum, 70.
sp., 70.
Zoo e c ie s, 27 n.
180
EXPLANATION OF THE PLATES.
PLATE I.
ENTOPROCTA.
Loxocalyx, Loxosoma.
(All the figures are magnified to the same scale:
scale, in y-, is indicated at the foot of the Plate.)
Zeiss, C. Obj., afterwards reduced 2/3. The
Fig.
i.
Fig.
2.
Fig.
3-
Fig.
4-
Fig.
5-
Fig.
6.
Fig.
• 7-
Fig.
8.
Fig.
9-
Fig.
IO.
Fig.
1 1.
Fig.
12.
Fig.
13-
Fig.
14.
Fig.
iS-
Fig.
16.
Fig.
'7-
Fig.
18.
Fig.
19.
Fig.
20.
Fig.
21.
Fig.
22.
Fig.
23'
Fig.
24.
Fig.
25.
Fig.
26,
Fig.
27.
Fig.
28,
Loxocalyx leptoclini Harmer (p. 8)
Loxosoma annulatum n. sp. (p. 11)
Loxosoma sluitcri n. sp. (p. 9).
Loxosoma velatum n. sp. (p. 1 3)
. — 402. D., Stat. 204. Femalc, with two embryos (cmb.).
. — 195. D.1, Stat. 274. Young individual.
— 247. E.2, Stat. 310. Older, with bud.
— 195. D.2, Stat. 274. Bud somewhat older.
„ „ „ . Bud still older.
— 24. E., Stat. 53. Young individual.
373. A.2., Stat. 105. Oral view.
- 24. D.2, Stat. 53. Male, young.
„ Male, younger.
> Male, old.
Loxosoma cirriferum n. sp. (p. 14). — 337. J.2, Stat. 71. Female, old: cmb., embryo.
v „ „ „ „ B Female, old: b., buds; cmb., embryo (on
slide 337. C.2, Retepord).
» Male, young.
Loxosoma circulare n. sp. (p. i6j. — 24. B.2, Stat. 53. Female, with two embryos (cmb.).
„ » 11 — v » » Male, with three buds.
— _ ,, Individual with three buds.
Loxosoma troglodytes n. sp. (p. 17). — 131. T.1, Stat. 164. Individual with two buds.
v „ „ — 142. D., „ „ Two individuals, seen in situ, in the
compensation-sac of Lepralia ccllcporoidcs, of which the operculum (op.) and the ascopore (asc.)
are shown (on slide 142 A., L. ccllcporoidcs).
Loxosoma pusillum n. sp. (p. 16). — 133. G., Stat. 164. Specimen with four buds.
vvv vvv Young individual.
Loxosoma loricatum n. sp. (p. 21). — 177. N.\ Stat. 257. Oral view.
„ Side view.
■ Aboral view.
„ „ „ „ „ Aboral view, with old bud.
B „ „ „ „ Individual, in situ, on Canda: se, scutum
of the Canda; set., its seta; Vib., its
vibraculum.
112.D.3, Stat. 156. Aboral view: b., bud; st., stalk.
B „ „ Oral view: st., stalk.
„ v v Oral view of an individual with embryos
(cmb.); st., stalk.
Fig. 29. Loxosoma breve n. sp. (p. 19). -- 131. U. Stat. 164. Two individuals, attached to the operculum
(op.) of Schizoporella. This and the next two
figures are fróm slide 131.B.3, Schizoporella.
„ „ „ — „ v v Another individual, on an operculum (op.).
„ „ v „ „ „ A larger specimen, on an operculum (0/.).
Loxosoma subsessile n. sp. (p. 19). -- 260. G., Stat. 318. This and the following figure are from
slide 260. D.1, Conescharellina.
, „ „ „ „ r Older individual, with two buds (^.) ; i7.,
stalk.
Loxosoma cocciformc n. sp. (p. 22).
Fig.
30.
Fig.
3i
Fig.
32
Fig.
33
Siboga- Expeditie. XXVI \\a, — S. F. Harmer, Polyzoa.
emb.
0
10
11
-emb.
emb.
17
m
*tar2\\
emb.
15
12
19
-b.
18
asc.
13
op
50
21
£&$V
27
f II
ÏU 32
Str''
op.
29
/
28
23
24
emb.
33
31
26
-b.
-Sfc.
op
~st.
st.
? r y r f r
1 1 mh il 1 1 1) imii II 1 1 1 umi
II I I 1 1 1 1 tl M I I M I ; ■ i ■ I ■ ;
S. V. Harmei del.
KXTOPKOCTA.
Fa. P. \V. M. Trap impr
PLATE II.
ENTOPROCTA.
Loxocalyx, Loxosoma, Pedicellina, Barentsia.
(All the figures are magnified to the same scale : — Zeiss, C. Obj., afterwards reduced '/i- The
scale, in /-:, is indicated at the foot of the Plate.)
Fig. i. Loxocalyx lineatus n. sp. (p. 6). — 332. C', Stat. 64. Aboral view.
Fig. 2. „ „ „ — „ „ „ Oral view.
Fig. 3. „ „ „ Oral view.
Fig. 4. Loxosoma lanchesteri n. sp. (p. 8). — Singapore, slide B (not 'Siboga'-Expedition). Aboral view.
v „ „ — Singapore, slide B (not 'Siboga'-Expedition). Calyx and part
of stalk, aboral view.
Barentsia geniculata n. sp. (p. 33). — 260. E., Stat. 318. Side view.
„ „ ■ „ Stalk-joint.
Barentsia discreta Busk (p. 29). — 195. B.', Stat. 274. Base of stalk, with stolons, one of which
is producing a young individual.
„ „ „ „ „ „ Old individual: an incipient joint is seen
at the middle of the stem; B, part of
body-wall of stalk.
Fig. 10. Barentsia laxa Kirkpatrick (p. 32). — 3S0. D.1, Stat. 47. Base (ö.) of stalk (st.), with appendages (app.).
Fig. 11. , „ „ — 88. A., Stat. 115. Old individual.
Fig. 12. Barentsia gracilis M. Sars (p. 27). — 262. F. Stat. 319. Side view of one individual.
Fig. 13. Pedicellina compacta n. sp. (p. 24). — 293. B.3, Stat. 273. Side view, with parts of stolons.
Fig. 14. „ „ „ — „ „ „ Young individual, with stolons.
Fig.
5.
Fig.
6,
Fig.
7-
Fig.
8.
Fig.
9
Siboga-Expeditie. XXVIIIa. — S. F. Harmer, Polyzoa.
//
S. F. Harmer del.
ENTOPROCTA.
Fa. P. W. M. Trap impr.
PLATE III.
CTENOSTOMATA.
Alcyonidium, Elzerina, Victorella, Arachnidiitm, Arachnoidea.
(Figs i and 10 were drawn vvith a Zeiss C Obj.; figs 2 — 9, ir, 14 and 15 with a Zeiss A Obj.;
and figs 12 and 13 with a Ross 2-inch Obj. All the figures were afterwards reduced '/.,. The scales, in /*,
are indicated at the foot of the Plate.)
Fig. 1. Alcyonidium polyoum Hassall (p. 37). — Torres Straits (not 'Siboga'-Expedition). Six zooecia,
growing on Euthyris obtecta, the zooecia of which are seen through the Alcyonidium.
Fig. 2. Elzerina blainvillii Lamx (p. 38). — Torres Straits (not 'Siboga'-Expedition). Distal end of a branch.
Arachnidium irregulare n. sp. (p. 48). — 376. B.4, Stat. 213. Zooecium.
„ „ „ „ „ „ Two zooecia, the lower one with the
kamptoderm everted.
„ „ „ „ „ „ Zooecium, with embryo in ovisac :
basal view.
„ „ „ — 376. B.2, „ v Another zooecium, with polypide :
basal view.
Arachnoidea protecta n. sp. (p. 50). — 378. E.2, Stat. 71. Zooecium with collar partially protruded.
„ n „ n „ „ Two zooecia in side view. Part of the
collar, in its retracted position, is seen
in the right hand zooecium.
„ „ „ — „ „ „ Zooecium with its collar protruded.
„ „ B „ „ „ Collar of the same specimen, more highly
magnified.
„ „ „ „ „ „ Another zooecium, with short peristome.
Victorella sibogae n. sp. (p. 45). — 451.B.2, Stat. 64. A group of old zooecia.
» v » v v A group of young zooecia.
„ „ „ i) » » An old zooecium, with degenerated polypide;
more highly magnified.
Fig. 15. „ „ „ „ „ „ An old zooecium, with functional polypide:
magnification as in fig. 14.
Fig.
3-
Fig.
4-
Fig.
5-
Fig.
6.
Fig.
7-
Fig.
8.
Fig.
9-
Fig.
10.
Fig.
1 1.
Fig.
12.
Fig.
13-
Fig.
14.
Siboga-Expeditie. XXVnfa. — S. F. Harmer, Polyzoa.
///
7)
P^N{
14
,,;-■
0 100 ZOO 300 «00 SOO
1 50 I I I I I
C.I I I I I I I I l I I
0 100 200 JOO 400 SOO 600 700 SOO 900 1000
A-ITll llllll I lllllllll I
O 1000 2000 3000 «000
2"\
S. F. Harmer del.
C rENOSTOMATA.
Fa. P. W. M. Trap impr.
PLATE IV.
CTENOSTOMATA.
Nolella (= Cylindroecium auctt.).
(Figs i — iS and 20 were drawn with a Zeiss A Obj., Fig. 19 with a Zeiss C Obj. All the figures
«rere afterwards reduced ' ... The scales, in ,a, are indicated at the foot of the Plate.)
Fig- 3
Fig. 4.
Fig.
5-
Fig.
6.
Fig.
7-
Fig.
8.
Fig.
9-
Fig.
10.
Fig. I. Nolella annectens n. sp. (p. 57). — 3S0. I.1, Stat. 47. A young and much elongated zooecium.
Fig. 2. „ „ „ » » i) Growing end of a branch. The dilated part
represents the commencement of a zooecium,
and already shows a young polypide-bud.
„ v „ „ „ v Zooecium with long peristome. The retracted
collar is seen in the vestibule.
v r „ r „ „ Zooecium with shorter peristome, showing
the polypide, the retracted collar and the
muscles.
v v „ r „ A similar specimen, in side view.
r v „ r „ „ A zooecium with shorter peristome, showing
the retracted collar.
A similar zooecium in side view.
T) ïï Tï V V 77
„ „ T v r „ A zooecium with very short peristome, in
side view.
B „ „ „ „ B Distal end of a zooecium with protruded collar.
Nolella papuensis Busk (p. 53). — 337. G. 2, Stat. 71. Zooecium with regenerated distal end, in
which are seen two embryos. An egg occurs
close to the distal end of the old part.
„ „ , T v v Part of a similar zooecium, containing distally
an embryo and, near the proximal end ot
the regenerated part, one or two eggs. The
outline is irregular in this region, indicating
that Iarvae have probably escaped here.
r „ „ r v v Distal end of a zooecium with three embryos.
„ v n v „ v Proximal end of a zooecium, showing
branches.
Stat. 164. Young zooecium, with branches.
Stat. 250. A long and slender zooecium.
Stat. 164. An elongated zooecium.
• 337. G.2, Stat. 71. A similar zooecium, showing, near the middle
of its length, two embryos which appear
to be about to escape.
r „ , An old and much shortened zooecium.
- 108. P. Stat. 144. A zooecium, more highly magnified, showing
several rounded bodies, possibly eggs, in
the vestibule.
„ „ „ Several zooecia, from the same slide, magnified
to the same extent as figs 1 — 18.
Fis. 11.
hig. 12.
Fig. 13-
Fig. 14.
Fig. 15.
Fig. 16.
Fig. 17.
130. H.3
276. A.,
130. H.3
Fig.
Fig.
18.
19.
(? sp.).
Fig. 20.
Siboga-Expeditie. XXVIII«. — S. F. Harmer, Polyzoa
i 100 200 300tOO S0060D 700 800 9001000
A-l"llllllllllllllllill
S. F. Harmer del.
CTKNOSTOMATA.
Fa. P. W. M. Trap impr.
PLATE V.
CTENOSTOMATA.
Amatliia, Bitskia.
(Figs i and 7 were drawn vvith a Ross 2-inch Obj.; figs 2 — 4, 6, 8, 12 and 14 with a Zeiss AObj.;
and figs 5, 9 — ir, 13 and 15 — 17 with a Zeiss C Obj. All the figures were afterwards reduced x\v The
scales, in ft, are indicated at the foot of the Plate.)
Amathia convoluta Lamx (p. 64). — 298. A.4, Stat. 273. Part of the zoarium.
— 2q8. A.5, , _ Zooecia in various conditions.
„ — 298. A.4, „ „ A single group of zooecia.
— 198. A.~', Stat. 274. Part of the stem, with attachments of
zooecia.
„ — 298. A.2, Stat. 273. Part of the stem, with young zooecia.
Amathia distans Busk (p. 68). -- 371. C.2, Stat. 273. Part of the stem, with a single group of
zooecia.
— „ , Part of zoarium, less highly magnified.
VVV VVV ' O J o
Buskia setigera Hincks (p. 87). — 339. B.2, Stat. 71. Group of zooecia.
v — 38. K.4, Stat. jj. Distal end of a zooecium, with protruded
collar (more highly magnified).
— „ _ Zooecium, attached to a Hydroid.
Buskia pilosa n. sp. (p. 89). — 501. E.-, Stat. 163. Part of stolon (st.), attached to the rootlets (r.)
of Caberea.
„ „ A zooecium, side view.
„ B „ A zooecium, more highly magnified, frontal
view.
307. B., Stat. 273. Part of zoarium.
• 451. B.', Stat. 64. Stolon and zooecium, on a zooecium of Victorclla
tig.
1.
Fig.
2.
Fig.
3-
Fig.
4-
Fig.
5-
Fig.
6.
Fig.
7-
Fig.
8.
Fig.
9-
Fig.
10.
Fig.
1 1.
Fig.
12.
Fig.
13-
Fig.
14.
Fig.
15-
Fig.
16.
Fig.
17-
Buskia nitcns Alder (p. 85)
ij » n
sibogae.
— 307. B., Stat. 273. Zooecium, side view.
— B „ Stolon and zooecium with protruded tentacles.
Siboga-Expedilie. XXVIIIV7. — S. F. Harmer, Polyzoa.
<
V
\
i
15
00 *O0 5>0
C-U-J- 1 I I I I I I l
0 100 ZOO 3ofl "90 ito 600 700 800 900 '000
A-l" -ihlilililili!
0 ,.« 1O0O 2000 30 00 4000 SOOO
2:'
I I I
m
I
12
V
16
17
S. F. Harmer del.
CTENOSTOMATA.
Fa. P. W. M. Trap impr.
PLATE VI.
CTENOSTOMATA.
Vesicularia, Valkeria.
(Figs I, 2, 8, 9, 13 — 15, 17 — 20 were dravvn with a Zeiss AObj.; figs 3 — 7, 10 — 12 and 16 with
a Zeiss C Obj. All the figures were aftervvards reduced 'ƒ2- The scales, in |t«, are indicated at the foot of
the Plate.)
Fig.
1.
Fig.
2#
Fig.
3-
Fig.
4-
Fig.
5-
Fig.
6.
Fig.
7-
Fig.
8.
Fig.
9-
Fig.
10.
Fig.
1 1.
Fig.
12.
Fig.
13-
Fig.
14.
Fig.
'5-
Fig.
16.
Fig.
>7-
Fig.
18.
Fig.
19.
Fig.
20.
Vesicularia papuensis Busk (p. 61). — 307. A.c, Stat. 27
I i-
Valkeria atlantica Busk (p. Ji). —
Valkeria tuberosa Heller (p. 76).
Part of the main stem, with branches.
„ „ „ Terminal branches, with zooecia.
„ „ „ Tentacles, protruded.
— „ „ „ Part of a branch, with two zooecia;
g., gizzard.
337. B.2, Stat. 71. Part of stolon, with zooecia, mostly old.
„ „ „ Another part of the stolon, with a young
(left) and an old (right) zooecium.
„ „ „ Group of zooecia.
B B „ Part of zoarium, less magnified.
393. C.1, Stat. 133. Part of zoarium, with zooecia elongated
at their proximal ends.
, v „ Zooecium of the same form.
„ „ „ Another zooecium of the same type.
v „ „ Part of stolon, with a similar zooecium.
130. G., Stat. 164. Old group of zooecia, drawn from an
unmounted specimen.
„ » b Young group of zooecia, unmounted.
Stolon from the basal side, showing the
mode of origin of a group of zooecia.
Two zooecia, more highly magnified.
Zooecium with kamptoderm and collar
protruded.
Distal end of zooecium with protruded
collar.
Distal end of zooecium with degenerated
polypide.
A similar zooecium, more contracted.
— 130- G.5,
— 130. G.2,
— 130. G.5,
Siboga- Expeditie. XXV1II«. — S. F. Harmer, Polyzoa.
VI
S. F. Harmer del.
0 100 200 300 400 500 «00 700 800 900 1000
CTENOSTOMATA.
Fa. P. W. M. Trap impr.
PLATE VIL
CTENOSTOMATA.
Bowerbankia, Mimosella.
(Figs I — 4, 6 — 9, il — 14 and 16 were drawn with a Zeiss C Obj.; figs 5, 10 and 15 were drawn
with a Zeiss A Obj. All the figures were afterwards reduced '/2- The scales, in ft, are indicated at the foot
of the Plate.)
Fig.
1.
Fig.
2,
Fig.
3<
Fig.
4
Fig.
5-
1. Mimosella bigeminata Waters (p. 79).
Fig.
Fig.
Fig.
Fig.
Fig.
12.
Fig.
13-
Fig.
14.
Fig.
iS-
Fig.
16.
38. M.4, Stat. Jj. Distal end of colony, with young zooecia.
„ „ „ v Part of the same, seen from the opposite
side.
„ „ „ B „ „ Part of the stem, with zooecia.
Three stem-internodes.
„ » » » » r> The greater part of a stem, with its
zooecia. 15 stem-internodes have been
omitted in the region just below the
persisting zooecia.
„ „ „ „ „ „ Part of the adnate stolon: /.', the primary
internode, giving off, on the opposite
side, s., the base of an erect stem ;
i.\ second internode of the main sto-
lon; Li.1, Li.1, successive internodes
of a lateral branch; r.1, main rootlet;
r.2, lateral rootlet.
„ B B — „ „ B single zooecium, showing the polypide.
Mimosella verticillata Heller (p. 81). — 139. D.2, Stat. 164. A single zooecium, with the base of
another.
„ „ „ , B „ Part of the adnate stolon, with the
primary internode (/.) and the bran-
ches to which it gives rise. The
proximal ends of five of the zooecia
are indicated.
v Part of the adnate stolon, with five
groups of zooecia.
Part of the adnate stolon, with two young
zooecia.
„ A single zooecium.
„ A single zooecium.
„ A single zooecium.
374. K. Loc. incert. Group of zooecia.
„ „ „ Portion of stem, with a zooe-
cium and parts of several others.
Fig. 10.
Fig. 11. Mimosella tennis n. sp. (p. 84). — 31. H.2, Stat. 53.
17 » 1) "'t' ^' ) V
Bowerbankia imbricata Adams (? sp.) (p. 70). —
Siboga-Expeditie. XXVIItrt. — S. F. Harmer, Polyzoa.
VII
S. F. Harmer del
CTENOSTOMATA.
Fa. P. W. M. Trap impi
PLATE VIII.
CYCLOSTOMATA.
Crisia.
(All the figures are magnified to the same scale :
The scale, in |Ct, is indicated at the foot of the Plate.)
Ross, i-inch Obj., afterwards reduced '/,
Fig.
i.
Fig.
2.
Fig.
3-
Fig.
4-
Fig.
5-
Fig.
6.
Fig.
7-
Fig.
8.
Fig.
9-
Fig.
10.
Fig.
1 1.
Fig.
12.
Fig.
'3-
Fig.
H-
Fig.
'5-
Fig.
i,6.
Fig.
'7-
Crisia elongata M.Edw. (p. 96). — 107. B.2, Stat. 144. Part of zoarium, with ovicell.
„ „ „ „ „ „ Part of branch, with rootlet.
» » » — 208. B.2, Stat. 282. Proximal internodes.
„ „ „ — 253. A.2, Stat. 315. Basal view of internodes, with ovicèll.
„ „ v — 107. B.2, Stat. 144. Proximal internodes.
„ „ T „ T v Distal view of internode, with ovicell,
showing ooeciopore.
„ r T ft „ ft Internodes, with ovicell.
„ , „ — 313. A., Stat. 273. Internodes, with twin-ovicells.
Crisia kerguelensis Busk (p. 105). — 37. S., Stat. jj. Proximal part of zoarium.
, „ „ — „ ft „ Distal part of zoarium.
„ „ „ — „ „ , Part of internode, with ovicell.
Crisia geniculata M.Edw. (p. 106). — Torres Straits, 88 (not 'Siboga'-Expedition). Part of zoarium.
Crisia cuneata Mapl. (p. 103). — 120. B., Stat. 164. Part of zoarium.
r „ „ — Japan, 5 AM. (not 'Siboga'-Expedition). Part of zoarium.
ft ft „ — ft „ (not 'Siboga'-Expedition). Internode, with ovicell.
, ft „ — ft , (not 'Siboga'-Expedition). Internode, with ovicell.
, „ „ — „ , (not 'Siboga'-Expedition). Internode, with ovicell.
Siboga-Expeditie. XXVIII<7. — S. F. Harmer, Polyzoa.
vin
S. F. Harmer del.
CYCLOSTOMATA.
Fa. P. W. M. Trap impi
PLATE IX.
CYCLOSTOMATA.
Tiibulipora.
(All the figures are magnified to the same scale : — Ross, i-inch Obj., afterwards reduced
The scale, in y., is indicated belovv the middle of the Plate.)
V.-
Fig. i. Tubulipora pulcherrima Kirkp. (p. 129). — ■ 92. H., Stat. 117. Part of a small zoarium, with ovicell:
<?., ooeciostome.
Fig. 2. „ „ „ — 350. C.2, Stat. 240. Branch with au ovicell hearing six
ooeciostomes (o.).
Fig. 3. „ „ „ — 99. A.1, Stat. 129. Fragment of a branch, with rooting
processes (r.).
Fig. 4. „ „ „ — » r> v Part of a reticulate colony, with two
ovicells: o., ooeciostomes.
Fig. 5. „ „ „ v v n Basal surface of two branches, showing
connecting kenozooecia (k.).
Fig. 6. Tubulipora cassiformis n. sp. (p. 135). — 1 12. A.1, Stat. 156. Parts of three branches, with con-
necting kenozooecia (£.); s., shield-
like portions of zooecia concealing
the orifices.
Fig. 7. „ r „ — 94. A.2, Stat. 119. Part of a colony, with ovicell and
connecting kenozooecia (/•.); o.,
ooeciostome.
Fig. 8. , „ „ „ „ „ Basal surface of three branches, with
connecting kenozooecia (£.).
Fig. 9. v „ „ — 1 12. A.1, Stat. 156. Side view of part of a branch, showing
the remarkable shields (s.) which
overlap the orifices (or.); k., keno-
zooecia of edge of branch.
Fig. 10. „ v „ — » ii » Basal view of branches, with con-
necting kenozooecia (k.).
Siboga-Expeditie. XXVIUrt. — S. F. Harmer, Polyzoa.
IX
S. F. Harmer del.
CYCLOSTOMATA.
1.1' W. M. Trap impr.
PLATE X.
CYCLOSTOMATA.
Entalopliora, Reptotubigera, Tubulipora, Crïsina.
(All the figures are magnified to the same scale : — Ross, i-inch Obj., afterwards reduced '/2.
The scale, in f4, is indicated at the foot of the Plate.)
Fig. i. Tubulipora atlantica (Forbes, MSS.), Johnst., var. fiexuosa, Pourt. (p. 127). — ni.D.1, Stat. 156.
Part of a branch, with ovicell.
Fig. 2. „ „ „ „ „ , „ — in. D.\ Stat. 156. Basal
view of branch.
Fig. 3. „ „ „ „ „ — iii.D.', Stat. 156. Branch,
with part of ovicell.
Fig. 4. Tubulipora atlantica (Forbes, MSS.), Johnst. (p. 124). — 165. B.1, Stat. 248. Basal view of branch,
showing secondary
thickening.
Fig. 5. „ „ Part of colony, with
ovicell : o., ooecio-
stome.
Fig. 6. Crisina radians Lamk (p. 139). — 336. A.2, Stat. 37. Branch with ovicell: ze/., porous lateral
windows of ovicell.
Fig. 7. „ „„ — „„„ Side view of fertile branch, showing zo.,
lateral windows of ovicell, and o., ooeci-
ostome.
Fig. 8. „ „„ — „ „ „ Basal view of branches.
Fig. 9. Reptotubigera philippsae n. sp. (p. 120). — 250. A.\ Stat. 310. Complete colony, with three ovicells:
/., marginal lamina with pore-areas; o., ooeciostomes.
Fig. 10. Tubulipora concinna MacGill. (p. 123). — 350. D., Stat. 240. Complete colony, with ovicells; o.,
ooeciostomes.
Fig. 11. Entalopliora delicatula Busk (p. 110). — 376. L., Stat. 213. Branches with ovicell: o., ooeciostome.
Fig. 12. Entalopliora proboscidea M.Edw. (p. 108). — 239. A.2, Stat. 310. Branches, with ovicell: o., ooeciostome.
Fig. 13. Entalopliora intricaria Busk (? sp.) (p. 112). — 135. F., Stat. 164. Branch without ovicell.
Fig. [4. „ , „ „ „ „ „ Branches, with ovicell: o., ? ooe-
ciostome.
Siboga- Expeditie. XXVIIT0. — S. F. Harmer, Polyzoa.
X
f 1
S. F. Harmei' del.
CYCLOSTOMATA.
Fa. P. W. M. Trap impr.
PLATE XI.
CYCLOSTOMATA.
Berenicea, Tervia, Hornera.
(Fig. 5 was drawn with a Zeiss A Obj., the other figures with a Ross i-inch Obj.; all the figures
were afterwards reduced 'L. The scales, in (i, are indicated near the middle and at the foot of the Plate.)
Fig. I. Tervia jellyae n. sp. (p. 143). — Queensland (not 'Siboga'-Expedition). Frontal view, showing the
distal end of au ovicell:
o., ooeciostome.
Fig. 2. „ ,, „ „ „ „ Basal viewofthesamebranch,
showing the ovicell.
Fig. 3. „ „ „ — 285. A., Stat. 51. Part of a branch, in frontal view.
Fig. 4. Berenicea sarniensis Norman (p. 1 14). ■ — 55 1. C, Stat. 310. Colony with three ovicells: o., ooeciostome.
Fig. 5. „ „ „ — „ „ „ Part of a younger colony, more highly
magnified.
Fig. 6. Berenicea lineata MacGill. (p. 116). — 104. C, Stat. 139. Young colony, with ovicell: o., ooeciostome.
Fig. 7. „ „ „ — 424. A., Stat. 240. Older colony, in which the ovicells form an
almost complete ring: o., ooeciostomes.
Fig. 8. Hornera spinigera Kirkpatrick (p. 147). — 112. B.', Stat. 156. Part ofa branch in which the orifices
are prolonged into specially long
spines.
Fig. 9. „ v „ — 82. A.2, Stat. 105. End of a young branch, from another
colony.
Fig. 10. „ „ „ — 1 12. B.1, Stat. 156. Branch with spinous zooecia.
Fig. il. „ „ „ — v „ „ Basal view of several branches, show-
ing an ovicell.
Fig. 12. „ „ „ — „ „ „ Frontal view of the same specimen:
o., ooeciostome.
Fig. 13. Hornera caespitosa Busk (p. 149). — 82. B., Stat. 105. Frontal view of several branches.
Siboga-Expeditie. XXVII.<ir. — S. F. Harmer, Polyzoa.
XI
r'Obj.
S. F. Haimer del.
CYCLOSTOMATA.
Fa. I' W. M. Trap impr.
Cf
PLATE XII.
CYCLOSTOMATA.
Super cytis, Liclienopora.
(Figs 2 and 7 were drawn with a Zeiss B Obj., the other figures with a Ross i-inch Obj.; all the
figures were afterwards reduced '/.2. The scales, in &. are indicated with figs 2 and 7 and at the foot of
the Plate.)
Fig. 1. Supercytis watersi n. sp. (p. 151). — 400. A., Stat. 164. Complete colony, showing the ovicell (ov.).
Fig. 2. Lic/wnopora mediterranen Blainv. (? sp.) (p. 164). — " 565. A., Stat. 125. Openings of cancelli, showing
the pin-like spines.
„ „ „ „ „ The complete colony from
which fig. 2 was drawn.
Lichenopora bnski n. sp. (p. 161). — 394. P., Stat. 144. Complete colony.
„ „ „ — Tosa, Japan (not'Siboga'-Expedition). Complete colony, showing
' the roof of the ovicell (ov.): o., ooeciostome.
Fig. 6. Liclienopora novae-zelandiae Busk (p. 155). — 202. B., Stat. 282. Colony with two young ovicells, the
roofs of which are not completely
formeel : o., ooeciostome.
„ „ r — „ „ „ Cancellus, with spines, more highly
magnified (from the colony shown
in fig. 8).
„ „ „ — „ „ n Colony with specially long peris-
tomes : o., ooeciostomes.
„ „ „ — 568. B.\ Stat. 105. Colony, with incompletely formed
or broken ovicell.
„ „ — 354. A., Stat. 240. Colony with short peristomes.
„ » » ■ 349- I-i Stat. 144. Colony with short peristomes. The
roof of the ovicell is not concealed
by cancelli : o., ooeciostome.
Fig.
3
Fig.
Fig.
4
5
Fig.
7-
Fig.
8.
Fig.
9-
Fig.
10.
Fig.
1 1.
Siboga- Expeditie. XXVIIIrt. — S. F. Harmer, Polyzoa.
XII
S. I'. Harmer del.
CYCLOSTOMATA.
Fa. I'. W. M. Trap impr.
Livr. LXXV.
RÈSULTATS DES EXPLORATIONS
ZOOLOGIQUES, BOTANIQUES, OCEANOGRAPHIQUES ET GEOLOGIQUES
ENTUEPRISES AUX
INDES NÉERI.ANDAISE3 OKIENTAI.ES Cn 1899 1900,
a bord du ÖIBOG-A
SOUS I.E COMMANDEMENT DE
G. F. TYDEMAN
PUDI.IÉS PAR
MA-X "W" E B E R.
Chef de 1'eipcdition.
*I.
«II.
•111.
IV.
*\\iis.
V.
• VI.
•VII.
• vin.
*IX.
*.\.
• XI.
•XII.
•XIII.
XIV.
*xv.
•XVI.
•XVII.
xvin.
XIX.
*xx.
•XXL
XXII.
XXIII.
*XXI\'i.
XXIV».
•Xxv.
•XXVI.
*XXVI4«.
XXVII.
•XXVIII.
•XXIX.
• XXX.
•XXXI.
•XXXII.
XXXIII.
•XXXIV.
XXXV.
•XXXVI.
• XXXVII.
XXXVIII.
• XXXIX.
■ *XL.
XLI.
• XLII.
• XLIII.
«XL1V.
*XLV.
XLVI.
• XLVII.
• XLYIII.
• XLIX'.
•XLIX2.
*L.
*LI.
•Lil.
•LUI.
•LIV.
LV.
• LVI.
*LV1I.
LVIII.
•LIX.
*LX.
•LXI.
• LXII.
LXIII.
LXIV.
LXV.
LXVI.
IutroJuction et description de l'expédition, Max Web er.
Le batcau et son équipement scientifique, G. F. Tv de man.
Résultats hydrographiques, G. F. Tydcman.
Foramiuifera, F. W. Winter.
Xenophyophoro, F. E. Sc hul ze.
Radiolaria, M. Hartmann.
Porifcra, G. C. J. Vosmaer et I. Ijiraa ').
Hydropolypi, A. Billard').
Stylasteriüa, S. .1. Hickson et MU« H. M. England.
Siphonophorn, Miles Lens et van Riemsdijk.
Ilydromedusae, O. Maas.
Seyphomcdiisae, O. Maas.
('tcnophora, MU« F. Moser.
Gorgouidae, Alcyouidae, J. Versluys, S. J. Hickson,
[O. C. Nut t ing et J. A. Thomson ').
Pennatulidae, S. J. Hickson.
Actiniaria, P. Mc Murrich1). .
Madreporaria, A. Alcock et L. Döderlein ').
Autipatharia, A. J. van Pesch.
Turbellaria, L. von Gr af f et R. R. von St umin er.
Cestodes, J. W. Spcugel.
Nematomorpha, H. F. Nievstrasz.
Chaetognatha, G. H. Fowler.
Ncmcrüni, A. A. W. Hub recht et Mllc G. Wijnhoff.
Myzostomidac, 11. R. von Stummer.
Polychacta errantia, R. Horst1).
Polychaeta sedentaria, M. Caullery et F. Mesnil.
Gephyrca, C. Ph. Sluiter.
Entcropneusta, J. W. Spengel.
Ptcrobranchia, S. F. H armer,
lirachiopoda, J. F. van Bemmelcn.
Polyzoa, S. F. II a r m e r ' ).
Cop'epoda, A. Scott1).
Ostracoda, G. W. Muller.
Cirripedia, P. P. C. Hoek.
Isopoda, H. F. Nierstrasz ').
Aniphipoda, Ch. Pérez.
Caprellidae, P. Maycr.
Stomatopoda, II. J. Hansen.
Cumacea, W. T. Cal man.
Schizopoda, II. J. Hansen.
Scrgestidae, II. J. Hansen.
Ilecapoda, J. G. de Man et J. E. W. Ihle •).
Pantopoda, J. C. C. Lorna n.
Halobatidae, J. Th. Oudemans.
Crinoidea, L. Döderlein et Austin H. Clark1).
Echinoidea, J. C. H. de. Me ij e re.
Holothurioidea, C. Ph. Sluiter.
Ophiuroidea, R. Kohier.
Asteroidea, L. Döderlein.
Solenogastres, H. E. Nierstrasz.
Chitonidae, H. F. Nierstrasz.
Prosobranchia, M. M. Schep man.
Prosobranchia parasitica, H. F. Nierstrasz et M. M.
Opisthobranchia, R. Bergh. [S-chepraau.
Heteropoda, J. J. Tesch.
Pteropoda, J. J. Tesch.
Lamellibranchiata, P. Pelseneer et Ph. Dautzenberg ').
Scaphopoda, M'lo M. Boissevain.
Cephalopoda, L. J o u b i n.
Tunicata, C. Ph. Sluiter et J. E. W. Ihle.
Pisces, Max Web er.
Cetacea, Max W e b e r.
Liste des algues, Mms A. Web er ').
Halimeda, M»e E. S. Barton. (Mme E. S. Gepp).
Corallinaceae, Mme A'. Web er et M. Foslie.
Codiaceae, A. et Mme E. S. Gepp.
Dinoflagellata. Coccosphaeridae, J. P. L o t s y.
Diatomaceae, J. P. Lotsy.
Deposita manna, O. B. Böggild.
Re'sultats géologiques, A. Wichmann.
Siboga-Expeditie
THE POLYZOA OF THE 10GA EXPEDITION
BY
SIDNEY F. HARMER, sc.d., i.r.s.
Keeper of Zoology in the Briti-.li Museum (Natural History), London
PART I
ENTOPROCTA, CTENOSTOMATA and
CYCLOSTOMATA
With 12 plates
Monographe XXVIII a of:
UITKOMSTEN OP ZOOLOGISCH,
BOTANISCH, OCEANOGRAPHISCH EN GEOLOGISCH GEBIED
verzameld in Neder la ndsch Oost-Indië 1899 — 1900
aan boord H. M. Siboga onder commando van
Luitenant ter zee ie kl. G. F. TYDEMAN
UITGEGEVEN DOOR
Dr. MAX WEBER
Prof. in Amsterdam, Leider der'Expeditie
(met medewerking van de Maatschappij ter bevordering van het Natuurkundig
Onderzoek der Nederlandsclie Koloniën)
BOEKHANDEL EN DRUKKERIJ
TOOHIIEEX
E. J. BRILL
LEIDEN
Publié Octobre 1915
* Les numéros avec un astérique ont déja paru ; ceux marqués 1) seulement en partie
Voor de uitgave van de resultaten der Siboga-Expeditie hebben
bijdragen beschikbaar gesteld:
De Maatschappij ter bevordering van het Natuurkundig Onderzoek der Nederlandsche
Koloniën.
Het Ministerie van Koloniën.
Het Ministerie van Binnenlandsche Zaken.
Het Koninklijk Zoologisch Genootschap > Natura Artis Magistra" te Amsterdam.
De > Oostersche Handel en Reederij" te Amsterdam.
De Heer B. H. de Waal, Oud-Consul-Generaal der Nederlanden te Kaapstad.
M. B. te Amsterdam.
The Elizabeth Thompson Science Fund.
Dr. J. G. de M. te Ierseke.
CONDITIONS GÉNÉRALES DE VENTE.
1°. L'ouvrage du „Siboga" se composera d'une série de monographies.
2°. Ces monographies paraitront au fur et a mesure qu'elles seront prêtes.
3°. Le prix de chaque, monographie sera différent, mais nous avons adopté comme base générale du prix de
vente: pour une feuille d'impression sans fig. flor. 0.15; pour une feuille avec fïg. flor. 0.20 a 0.25;
pour une planche noire flor. 0.25 ; pour une planche coloriée flor. 0.40; pour une photogravure flor. 0.60.
4°. Il y aura deux modes de souscription :
a. La souscription a l'ouvrage complet.
b. La souscription a des monographies scparées en nombre restreint.
Dans ce dernier cas, le prix des monographies sera majoré de 25 °/0.
50. L'ouvrage sera réuni en volumes avec titres et index. Les souscripteurs a l'ouvrage complet recevront
ces titres et index, au fur et a mesure que chaque volume sera complet.
Déia paru: Prix:
J r Souscription Monographies
al'ouvrage complet séparées
ie Livr. (Monogr. XLIV) C. Ph. Sluiter. Die Holothurien der Siboga-Expedition. Mit toTafeln. / 6. — / 7.50
2e Livr. (Monogr. LX) E. S. Barton. The genus Halimeda. With 4 plates „ 1.80 „ 2.40
3e Livr. (Monogr. I) Max Weber. Introduction et description de 1'expédition. Avec Liste des
Stations et 2 cartes n 6.75 , 9. —
4e Livr. (Monogr. II) G. F. Tydeman. Description of the ship and appliances used for scientific
exploration. With 3 plates and illustrations „ 2. — ■ „ 2.50
5e Livr. (Monogr. XLVII) H. F. Nierstrasz. The Solenogastres of the Siboga-Exp. With 6 plates. „ 3.90 „" 4.90
6e Livr. (Monogr. XIII) J. Versluys. Die Gorgoniden der Siboga-Expedition.
I. Die Chrysogorgiidae. Mit 170 Figuren im Text .-...„ 3.— „ 3.75
7e Livr. (Monogr. XVI a) A. Alcock. Report on the Deep-Sea Madreporaria of the Siboga-
Expedition. With 5 plates. . ■. „ 4.60 „ 5.75
3e Livr. (Monogr. XXV) C. Ph. Sluiter. Die Sipunculiden und Echiuriden der Siboga-Exp.
Mit 4 Tafeln und 3 Figuren im Text „ 3. — n 3.75
9e Livr. (Monogr. Vla) G. C. J. Vosmaer and J. H. Vemhout. The Porifera of the Siboga-
Expedition. I. The genus Placospongia. With 5 plates „ 2.40 „ 3. —
lOe Livr. (Monogr. XI) OttO Maas. Die Scyphomedusen der Siboga-Expedition. Mit 12 Tafeln. „ 7.50 ,, 9.50
iie Livr. (Monogr. XII) Fanny Moser. Die Ctenophoren der Siboga-Expedition. Mit 4 Tafeln. „ 2.80 „ 3.50
12e Livr. (Monogr. XXXIV) P. Mayer. Die Caprellidae der Siboga-Expedition. Mit 10 Tafeln. „ 7.80 „ 9.75
13e Livr. (Monogr. III) G. F. Tydeman. Hydrographic results of the Siboga-Expedition. With
24 charts and plans and 3 charts of depths „ 9. — n 11.25
14e Livr. (Monogr. XLIII) J. C. H. de Meijere. Die Echinoidea der Siboga-Exp. Mit 23 Tafeln. „ 15. — „ 18.75
15e Livr. (Monogr. XLVa) René Koehler. Ophiures de 1'Expédition du Siboga.
ie Partie. Ophiures de Mer profonde. Avec 36 Planches „ 16.50 „ 20.50
i6e Livr. (Monogr. LII) J. J. Tesch. The Thecosomata and Gymnosomata of the Siboga-
Expedition. With 6 plates ■ „ 3.75 n 4.70
17e Livr. (Monogr. LVla) C. Ph. Sluiter. Die Tunicaten der Siboga-Expedition.
I. Abteilung. Die socialen und holosomen Ascidien. Mit 15 Tafeln „ 6.75 „ 9. — -
iSe Livr. (Monogr. LXI) A. Weber— van Bosse and M. Foslie. The Corallinaceae of the Siboga- '
Expedition. With 16 plates and 34 textfigures „ 12.50 n 15-50
19e Livr. (Monogr. VIII) Sydney J. Hickson and Helen M. England. The Stylasterina of
the Siboga Expedition. With 3 plates „ 1.50 „ 1.90
20e Livr. (Monogr. XLVIII) H. F. Nierstrasz. Die Chitoncn der Siboga-Exp. Mit 8 Tafeln. „ 5.— „ 6.25
2 ie Livr. (Monogr. XLV£) René Koehler. Ophiures de 1'Expédition du Siboga.
2e Partie. Ophiures littorales. Avec 18 Planches „ 10.25 n 12.75
22e Livr. (Monogr. XXVIfc) Sidney F. Harmer. The Pterobranchia of the Siboga-Expedition,
with an account of other species. With 14 plates and 2 text-figures „ 6.75 „ 9. —
23e Livr. (Monogr. XXXVI) W. T. Calman. The Cumacea of the Siboga Expedition. With
2 plates and 4 text-figures v 1.80 „ 2.40
24e Livr. (Monogr. LVLz) C. Ph. Sluiter. Die Tunicaten der Siboga-Expedition.
Supplement zu der I. Abteilung. Die socialen und holosomen Ascidien. Mit 1 Tafel. „ — .75 „ 1. —
25e Livr. (Monogr. L) Rud. Bergh. Die Opisthobranchiata der Siboga-Exped. Mit 20 Tafeln. „ 11.25 „ 14.10
26e Livr. (Monogr. X) OttO Maas. Die Craspedoteii Medusen der Siboga-Exp. Mit 14 Tafeln. n 9.25 B 12.50
27e Livr. (Monogr. XIII a) J. Versluys. Die Gorgoniden der Siboga-Expedition.
II. Die Prhnnoidae. Mit 10 Tafeln, 178 Figuren im Text und einer Karte . . . „ 12.50 „ 16.75
28e Livr. (Monogr. XXI) G. Herbert Fowler. The Chaetognatha of the Siboga Expedition.
With 3 plates and 6 charts n 4.20 „ 5.25
29e Livr. (Monogr. LI) J. J. Tesch. Die Heteropoden der Siboga-Expedition. Mit 14 Tafeln. n 6.75 „ 9.—
30e Livr. (Monogr. XXX) G. W. Muller. Die Ostracoden der Siboga-Exped. Mit 9 Tafeln. „ 3.50 „ 4.40
31e Livr. (Monogr. IV bis) Franz Eilhard Schulze. Die Xenophyophoren der Siboga-Exped.
Mit 3 Tafeln „ 2.40 „ 3.—
32e Livr. (Monogr. LIV) Maria Boissevain. The Scaphopoda of the Siboga Expedition.
With 6 plates and 39 textfigures B 4.80 6.
33e Livr. (Monogr. XXVI) J. W. Spengel. Studiën über die Enteropneusten der Siboga-Exp.
Mit 17 Tafeln und 20 Figuren im Text n 14. — v 17.50
34e Livr. (Monogr. XX) H. F. Nierstrasz. Die Nematomorpha der Siboga-Exp. Mit 3 Tafeln. „ 2.80 " 3.50
35e Livr. (Monogr. XIIL) Sydney J. Hickson und J. Versluys. Die Alcyoniden der Siboga-
Exped. I. Coralliidae, II. Pscudocladochonus Hicksoni. Mit 3 Tafeln und 16 Figuren im Text. v 2.20 „ 2.75
36e Livr. (Monogr. XXXIrt) P. P. C. Hoek. The Cirripedia of the Siboga Expedition.
A. Cirripedia pedunculata. With 10 plates c.40 6.75
37e Livr. (Monogr. Xhlla) L. Döderlein. Die gestielten Crinoiden der Siboga-Expedition. Mit
23 Tafeln und 12 Figuren im Text 8.— 10.
38e Livr. (Monogr. IX) Albertine D. Lens and Thea van Riemsdijk. The Siphonophores
of the Siboga Expedition. With 24 plates and 52 textfigures n 13.50 B 16.75
39e Livr. (Monogr. XLIX'rt) M. M. Schepman. The Prosobranchia of the Siboga Expedition.
Part I. Rhipidoglossa and Docoglossa, with an Appendix by Prof. R. Bergh.
With 9 plates and 3 textfigures 4.80 „ 6. —
40e Livr. (Monogr. XL) J. C. C. Loman. Die Pantopoden der Siboga-Expedition. Mit 15
Tafeln und 4 Figuren im Text B 6.25 „ 7.80
Prix:
Souscription Monogrnphies
a 1'ouvrage complet séparces
41e Livr. (Monogr. LVIr) J. E. W. Ihle. Die Appendicularien der Siboga-Expedition. Mit 4
Tafeln und 10 Figuren im Text / 4.80 ƒ 6. —
42c Livr. (Monogr. XLIX-) M. M. Schepman und H. F. Nierstrasz. Parasitische Proso-
branchier der Siboga-Expedition. Mit 2 Tafeln „ 1.20 „ 1.50
43c Livr. (Monogr. Xl.iX'A) M. M. Schepman. The Prosobranchia of the Siboga Expedition.
Part II. Taenioglossa and Ptenogiossa. With 7 plates ■ . . . . „ 4.50 „ 5.60
44c Livr. (Monogr. XXIX«) Andrew Scott. The Copepoda of the Siboga Expedition.
Part I. Ffeë-swimming, Littoral and Semi-parasitic Copepoda. With 69 plates. . „ 26. — „ 32.50
45c Livr. (Monogr. LVI£) C. Ph. Sluiter. Die Tunicaten der Siboga-Expedition.
II. Abteilung. Die Merosomen Ascidicn. Mit 8 Tafeln und 2 Figuren im Text. „ 5.75 „ 7.25
46e Livr. (Monogr^ XLIX'<j M. M. Schepman. The Prosobranchia of the Siboga Expedition.
Part III. Gymnoglossa. With 1 plate „ — .80 „ 1. —
47c Livr. (Monogr. XIIIZ>) C. C. Nutting. The Gorgonacea of the Siboga Expedition.
III. The Muriceidae. With 22 plates „ 8.50 „ 10.75
48e Livr. (Monogr. XIII^') C. C. Nutting. The Gorgonacea of the Siboga Expedition.
IV. The Plexauridae. With 4 plates „ 1.60 „ 2.—
49e Livr. (Monogr. LVIrt') J. E. W. Ihle. Die Thaliaceen (einschliesslich Pyrosomen) der
Siboga-Expedition. Mit 1 Tafel und 6 Figuren im Text. . . . „ 1.75 , 2. 20
50c Livr. (Monogr. XIII£2) C. C. Nutting. The Gorgonacea of the Siboga Expedition.
V. The Isidse. With 6 plates „ 2.25 „ 3.—
51e Livr. (Monogr. XXXVII) H. J. Hansen. The Schizopoda of the Siboga Expedition. With
16 plates and 3 text figures - . „ 12.75 „ 16. —
52e Livr. (Monogr. XlllP) C. C. Nutting. The Gorgonacea of the Siboga Expedition.
VI. The Gorgonellidae. With 11 plates „ 4. — „ 5. —
53c Livr. (Monogr. XV a) J. Playfair Mc Murrich. The Actiniaria of the Siboga Expedition.
Part I. Ceriantharia. With 1 plate and 14 text figures „ 2.20 „ 2.75
54e Livr. (Monogr. XIII^4) C. C. Nutting. The Gorgonacea of the Siboga Expedition.
VII. The Gorgonidae. With 3 plates „ 1.20 t 1.50
55e Livr. (Monogr. XXXIX a) J. G. de Man. The Decapoda 'of the Siboga Expedition.
Part I. Family Penaeidae „ 2.60 „ 3.25
56c Livr. (Monogr. LXII) A. & E. S. Gepp. The Codiaceae of the Siboga Expedition including
a Monograph of Flabellarieae and Udoteae. With 22 plates „ 12.50 „ 15.50
57e Livr. (Monogr. X\IIbr' C. C. -Nutting. The Gorgonacea of the-Siboga Expedition.
VIII. The Scleraxonia. With 12 plates. . . . . „ 4.80 „ 6. —
58e Livr. (Monogr. XLIX'a') M. M. Schepman. The Prosobranchia of the Siboga Expedition.
Part IV. Rachiglossa. With 7 plates „ 5.— , 6.25
59e Livr. (Monogr. VI«') G. C. J. Vosmaer. The Porifera of the Siboga-Expedition.
II. The genus Spirastrella. With 14 plates „ 6.20 „ 7.75
60e Livr. (Monogr. XXXIX ax) J. G. de Man. The Decapoda of the Siboga Expedition.
Part II. Family Alpheidae „ 6.40 „ 8. —
6i« Livr. (Monogr. LUIa) Paul Pelseneer. Les Lamellibranches de 1'Expédition du Siboga.
Partie Anatomique. Avec 26 planches „ IO. — v 12.50
62e Livr. (Monogr. XXIV' a) R. Horst. Polychaeta er-rantia of the Siboga Expedition.
Part I. Amphinomidae. With 10 plates „ 3.85 „ 4.80
63e Livr. (Monogr. Llllb) Ph. Dautzenberg et A. Bavay. Les Lamellibranches de 1'Expéd.
du Siboga. Partie Systématique. I. Pectinidés. Avec 2 planches „ 2.25 B 2.80
64e Livr. (Monogr. XLIX1^) M. M. Schepman. The Prosobranchia of the Siboga Expedition.
Part V. Toxoglossa. With 6 plates and 1 textfigure „ 4.80 „ 6. —
65e Livr. (Monogr. LVII) Max Weber. Die Fische der Siboga-Expedition. Mit 12 Tafeln und
123 Figuren im Text . * „ 22. — „ 27.50
66e Livr. (Monogr. XLIX/) M. M." Schepman. The Prosobranchia, Pulmonata and Opistho-
branchia Tectibranchiata Tribe Bullomorpha of the Siboga Expedition.
Part VI. Pulmonata and Opisthobranchia Tectibranchiata Tribe Bullomorpha. With
2 plates „ I.75 „ 2.20
67? Livr. (Monogr. XXXI ó) P. P. C. Hoek. The Cirripedia of the Siboga-Expedition.
B. Cirripedia sessilia. With 17 plates and 2 textfigures „ 8. — „ 10. —
68e Livr. (Monogr. hlXa) A. Weber-van Bosse. Liste des Algues du Siboga.
I. Myxophyceae, Chlorophyceae, Phaeophyceae avec Ie concours de M. Th. Reinbold.
Avec 5 planches et 52 figures dans Ie texte B 6. — „ 7.50
69e Livr. (Monogr. XXXIX«) J. G. de Man. The Decapoda of the Siboga Expedition.
Supplement to Part I. Family Penaeidae. Explanation of Plates „ 3.20 „ 4. —
70c Livr. (Monogr. VII a) A. Billard. Les Hydroïdes de 1'Expédition du Siboga.
I. Plumularidae. Avec 6 planches et 96 figures dans Ie texte . ; „ 5-75 » 7-2°
71e Livr. (Monogr. XXXIX7;) J. E. W. Ihle. Die Decapoda brachyura der Siboga-Expedition.
I. Dromiacea. Mit 4 Tafeln und 38 Figuren im Text „ 3-5° „ 4-4°
72c Livr. (Monogr. XXXII a) H. F. Nierstrasz. Die Isopoden der Siboga-Expedition.
I. Isopoda chelifera. Mit 3 Tafeln 1, "2.15 i 2-6°
73c Livr. (Monogr. XVII) A. J. Van PeSCh. The Antipatharia of the Siboga Expedition. With
8 plates and 262 textfigures „ 10.75 „ 13.50
74e Livr. (Monogr. XXXIX,?1) J. G. de Man. The Decapoda of the Siboga Expedition.
Supplement to Part II. Family Alpheidae. Explanation of Plates „ 7. — „ 8.75
75e Livr. (Monogr. XXVIII a) Sidney F. Harmer. The Polyzoa of the Siboga Expedition.
Part I. Entoprocta, Ctenostomata and Cyclostomata. With 12 plates „ 8.80 „-11. —
1 flor. ss Mrk 1.70 = I sli. 8 d. = frs 2.12 en chiflïes airondies.
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